Reviewing the benefits of marine sanctuaries by Save Our Marine Life

Reviewing the benefits of marine sanctuaries Asia Armstrong (BMarSt Hons Class I)

This report is an independent research paper commission by the Australian Marine Conservation Society

EXECUTIVE SUMMARY This review of more than 100 recent Australian and international research papers has identified there are many benefits that can flow from the creation of marine sanctuaries. Along with describing these benefits, the review also reports on public opinion surveys that demonstrate widespread community support for marine sanctuaries.

Marine sanctuaries may also be referred to as marine national parks, green zones or no-take zones and are protected areas free of extractive commercial and recreational activities. Although their primary objective is to conserve marine life and promote education and recreation, they can also provide benefits to the fishing and tourism sectors.

Sanctuaries for conserving biodiversity Extensive research has shown that well-managed sanctuaries of any size have larger and far more abundant marine life than nearby fished areas, as shown by studies of Mud Crab in NSW, Flathead in the Baltic Sea, Striped Marlin off California and lobsters in New Zealand. These differences become even greater with increases in a sanctuaryâ&#x20AC;&#x2122;s size and age, along with improvements in its management effectiveness.

The monitoring of marine sanctuaries reveals that they can have twice as many large fish species, five times more fish biomass and 14 times more shark biomass than nearby fished areas.

Although small sanctuaries are effective in protecting endangered animals that largely stay put, such as Dusky Groupers in the Azores, networks of large sanctuaries benefit migrating whales, sharks and seabirds by protecting their critical breeding and feeding habitats from human impact. Shark numbers in protected areas on the Great Barrier Reef are, for example, ten times that of unfished areas. Larger sanctuaries also include more habitats and help achieve global marine conservation targets.

Once established, sanctuaries encourage a return to more natural communities because fishing pressure has been removed. Research in NSW, Mexico and Florida has revealed that sanctuaries can lead to an increase in the number of predator species and adult spawning fish. This increases the areaâ&#x20AC;&#x2122;s natural productivity, in some places up to four times, and helps it

resist climate change and recover from flood damage and pest invasionsâ&#x20AC;&#x201D;crown-of-thorns starfish outbreaks are less frequent in sanctuary zones on the Great Barrier Reef.

Sanctuaries can also be used to reduce the number of cumulative impacts in a marine area, protect threatened species and generally improve the areaâ&#x20AC;&#x2122;s overall health, as shown by research in Tasmania, the Adriatic Sea and Brazil.

Benefits for fisheries There is considerable research to show that marine sanctuaries can benefit fisheries. The benefits can include increased commercial and recreation catches along sanctuary boundaries, such as in South Africa and St Lucia, as well as fish and larval spillover underpinning the sustainability of surrounding fisheries in 12 of 14 sanctuaries analysed.

The review also reports on the debate between the advocates of fisheries management and those of marine sanctuaries with regards to spillover and the recovery of marine life. The research indicates that fisheries management tools can be effective in increasing the catch of targeted fish; however they cannot match the performance of marine sanctuaries in the protection of non-targeted marine life. This is because the focus of fisheries management is largely on single species and not the wider ecosystem.

The integration of large marine sanctuaries and fisheries management tools will likely ensure the best environmental, social and economic outcomes, while the cost of managing sanctuaries that support fisheries is also much lower than fishery subsidies.

Socio-economic considerations There are many Australian and overseas studies indicating that marine sanctuaries boost the tourism industry. For example, NSW sanctuaries are assisting the recovery of Australiaâ&#x20AC;&#x2122;s east coast grey nurse shark numbers as well as providing tourism income to local dive operators whose dive trips have increased. Shark diving tourism in Palau is worth $18million each year, far higher than the $10,800 that would be generated by harvesting the local shark population.

Sanctuaries also provide important baseline sites for scientists to compare them with fished areas and so measure the impact of human activities on marine life as is being done in New Zealand, the Pacific and elsewhere.

Community Support for sanctuaries In general, the community endorses marine sanctuaries as important tools for biodiversity conservation, as shown by polling in Australia, the UK and New Zealand. A global survey has also revealed general acceptance of sanctuaries from commercial fishers, while polls in the Great Barrier Reef demonstrate significant support from recreational fishers and high levels of compliance with sanctuary rules.

Given the 2014 target set by the World Parks Congress to protect 30% of the global marine environment with sanctuaries by 2030, and given the multitude of benefits associated with these sanctuaries, retaining the current National Representative System of Marine Protected Areas in Australia would be a defensible and wise decision.

Reviewing the benefits of marine sanctuaries Asia Armstrong (BMarSt Hons Class I) - Asia Armstrong recently graduated from The University of Queensland with a Bachelor of Marine Studies, Honours Class I. Asia has experience in undertaking scientific research and literature reviews and continues her work at the University of Queensland with Project Manta, a collaborative initiative that researches the biology and ecology of manta rays in Australian waters. This report is an independent research paper commission by the Australian Marine Conservation Society.

CONTENTS Introduction

Benefits of marine sanctuaries

Sanctuaries for conserving biodiversity

Density, biomass and size

Community and population structure

Benefits beyond fish

Habitat protection and ecosystem resilience

Age and size do matter

Benefits to fisheries

Spillover effects and larval recruitment

Debate over sanctuaries

Socio-economic considerations

Tourism income

Benefit to science

Economic management

Community support for sanctuaries Compliance

14 15

Conclusion

References

INTRODUCTION Protected areas are recognized as central to conservation efforts for protecting land and marine resources. Protected Areas are officially defined by the International Union for Conservation of Nature (IUCN) as: “A clearly defined geographical space, recognised, 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). Six categories of protection have been defined by the IUCN, with each category distinguished by different levels of protection and prescribed objectives. National parks, whether terrestrial or marine, are defined as Category II protected areas. They “are large natural or near natural areas set aside to protect large-scale ecological processes, along with the complement of species and ecosystems characteristic of the area, which also provide a foundation for environmentally and culturally compatible spiritual, scientific, educational, recreational and visitor opportunities” (Dudley 2008). In the marine environment these areas are also referred to as marine national parks, marine sanctuaries, green zones or no-take zones. These areas form the focus of this current synthesis and henceforth will be referred to as marine sanctuaries.

The primary objective of marine sanctuaries is “to protect natural biodiversity along with its underlying ecological structure and supporting environmental processes, and to promote education and recreation” (Dudley 2008). In short, sanctuaries aim to conserve biodiversity and promote the human experience of nature in a non-extractive manner. However, complimentary benefits derived from their establishment include resilience to climate change, improvements to fisheries and socio-economic benefits. Despite the wealth of scientific evidence in favour of sanctuaries, there continues to be debate around their effectiveness. This report aims to provide a global summary of the most recent research surrounding the benefits of sanctuaries. The major focus is on biodiversity conservation, with attention also given to the benefits to fisheries, socio-economic considerations, and community support for sanctuaries.

BENEFITS OF MARINE SANCTUARIES Multiple reviews of the literature surrounding sanctuaries have been undertaken, with the overwhelming conclusion that well managed sanctuaries of any size harbour more diversity, higher abundance, and organisms of larger size, than adjacent fished areas (Gell & Roberts 2003, Halpern 2003, Lester et al. 2009, Costello 2014, Edgar et al. 2014, Watson et al. 2014).

Further, the conservation benefits of sanctuaries increase exponentially with the accumulation of 5 key features: no-take, well enforced, old (>10yrs), large (>100km2) and isolation by deep water or sand (Edgar et al. 2014). In addition, a global review of sanctuaries has determined that the following five variables were not significant predictors of overall sanctuary benefits: (1) location; (2) displaced fishing effort; (3) latitude; (4) size; and (5) taxonomic composition (Lester et al. 2009).

Sanctuaries for conserving biodiversity A worldwide investigation of 87 sanctuaries has shown that effective sanctuaries have twice as many large fish species (>250mm total length), five times more large fish biomass, and 14 times more shark biomass, than comparable fished areas (Edgar et al. 2014). Further, by using four biological measures of success, a global meta-analysis of 89 sanctuary studies showed higher values for sanctuaries compared to fished areas outside sanctuary zones (Halpern 2003). Additionally, scores increased after sanctuary formation compared to before, emphasising that effects are not simply an artefact of sanctuary location.

Density, biomass and size The benefit of sanctuaries to the biomass, size and abundance of fish assemblages is well supported in the literature. Examples are varied and many: the density, mean length and biomass of coral trout were consistently greater in marine sanctuaries than on fished reefs of the Great Barrier Reef over both short and medium term (Emslie et al 2015); increase in mud crab numbers in New South Wales (NSW) (Butcher et al. 2014); increases in mean nekton size in eastern Australia (Pillans et al. 2007); increased density of flathead in the Baltic Sea (Florin et al. 2013); 463% increase in biomass of fish species in Mexico (Aburto-Oropeza et al. 2011); increased abundance of striped marlin in California (Jensen et al. 2010); 80% higher biomass of limpets in South Africa (Branch & Odendaal 2003); 500% increase in catch per unit effort biomass for spiny lobsters in the Mediterranean over a 10 year period (Follesa et al. 2011); two times greater abundance and four times greater biomass of sharks in Fiji (Goetze & Fullwood 2013); increases in lobster biomass and egg production in New Zealand (Kelly et al. 2000); and higher relative abundance and mean length of four slowgrowing, high trophic level species from the Indian Ocean (Maggs et al. 2013). Finally, on a global scale, well managed sanctuaries contain more than five times the total large fish biomass and 14 times the shark biomass compared to fished areas (Watson et al. 2014).

Community and population structure Sanctuaries also impact community and population structures of the ecosystem. As is the situation in nature, some species benefit from protection at the detriment of others. In NSW sanctuaries, predators of seahorses (3 species of fish and 2 species of octopus) were higher than in fished areas, and seahorse numbers were negatively correlated with predator numbers (Harasti et al. 2014). Further, research into temperate reefs in Tasmania has shown that sanctuaries increase the number of predators such as lobsters, which results in lower numbers of urchins. This increases the resilience of kelp beds, strengthening the adaptive capacity of these ecosystems (Ling et al. 2009). Thus, sanctuaries promote a more natural complement of large and functional predators and minimize the risk of local phase shifts by reinstating habitat-specific predator-prey dynamics eroded by fishing (Ling & Johnson 2012). Additionally, sanctuaries in Mexico show predators exhibit 11 times higher biomass and carnivores four times higher biomass after protection than before protection (Aburto-Oropeza et al. 2011). This highlights the fact that protection can result in both winners and losers.

However, there are also demographic benefits that come with protection. In Florida, a number of exploited fish species showed increases in size, adult abundance, adult spawning and occupancy rates in response to closure from fishing (Ault et al. 2013). Sanctuaries such as those in western USA can protect older females, with high-quality larvae and can protect their range from overfishing the otherwise reproductively isolated populations (Berkeley et al. 2004). Further, Baltic Sea flathead show higher numbers of older individuals and more even sex ratios in response to protection (Florin et al. 2013). In South Africa, limpets in a sanctuary zone were found to be 65% larger, had 40% higher proportion of females, 25% greater survivorship and 33% greater growth. Wave-exposed sites within sanctuaries also increased recruitment of limpets by 45% compared to sites outside sanctuaries (Branch & Odendaal 2003).

Benefits beyond fish Sanctuaries are designed to protect a wide range of biodiversity that utilize the marine environment, including birds, sharks, mammals and their habitats. Sanctuaries benefit large marine mega-fauna, such as cetaceans (whales and dolphins), pinnipeds (seals and sea lions), sea otters, polar bears, seabirds, sharks, cephalopods and predatory fishes (Hooker & Gerber

2004). In turn, the presence of mega-fauna can provide important proxies for designating sanctuary zones. In the Great Barrier Reef (GBR), an order of magnitude (10 times) more sharks are found on reefs protected from fishing pressure (Robbins et al. 2005). Further, sanctuaries provide conservation benefits that cover a variety of shark species and their life history stages (Knip et al. 2012). Even highly mobile mega-fauna species benefit from networks of sanctuaries. In Canada, killer whales show a strong preference for sanctuary zones compared to adjacent waters, and they use them preferentially to feed and for beachrubbing (for parasite removal or social interaction) (Williams et al. 2009b). Sanctuaries comprise 0.001% of the killer whales range, but are utilized 6.5% of the time. In Greece, 72% of all turtle GPS locations were recorded within sanctuary zones, and these areas cover the majority of their breeding and foraging activity, highlighting their importance for this species conservation (Schofield et al. 2013). Sanctuaries in Brazil preserve mangrove estuaries and coral reef habitats and thus contain higher densities of manatees, dolphins and turtles that utilize these habitats, providing protection for these highly threatened species (Alves et al. 2013). Trophic cascades, whereby negative impacts at one level of a food web have knock on effects to other trophic levels, have been well documented globally. A recent study revealed that the loss of sharks from an ecosystem can have an impact that propagates down the food chain, potentially contributing to meso-predator release and altering the numbers of primary consumers (Ruppert et al. 2013). This same study revealed that bottom-up processes of bleaching and cyclones can propagate up the food chain through herbivores, planktivores and corallivores. Importantly, sanctuaries have been shown to protect both benthic and pelagic species, even those that move in and out of sanctuaries, and thus can help prevent both topdown and bottom-up trophic cascades (Costello 2014).

Despite the highly mobile nature of many bird species, sanctuaries can provide important protection for critical habitats that cover important life history stages for this group of animals. The Macquarie Island Marine Park provides protection to Southern and Northern giant petrels during breeding season, chick rearing and fledgling stages (Trebilco et al. 2008). However, the terrestrial habitat utilised by juveniles remains a substantial risk to these species, as this area remains unprotected. This highlights the need for complimentary marine and terrestrial protected areas that cover important life history stages of these species. Coastal sanctuaries in the Mediterranean overlap across multiple life stages for Yelkouan shearwaters, with 31% of foraging area, 38% of diving positions and 27% of resting positions protected, highlighting the importance of these areas for conservation of this species (PĂŠron

et al. 2013). The majority of the foraging grounds for African penguins, Bank cormorants and Cape gannets were covered by the boundaries of sanctuaries in Namibia, providing protection for a vital activity in these species life histories (Ludynia et al. 2012). Immediately following the establishment of a 20km area sanctuary in South Africa, resident African penguins reduced their foraging to within the protected area. At the same time, the foraging range expanded in a nearby colony that did not have access to the protected area, indicating an energetic cost associated with fisheries competition (Pichegru et al. 2010). However, when this was assessed after two years of implementation it was found that â&#x20AC;&#x153;fishing the lineâ&#x20AC;? (whereby fishing vessels concentrate their efforts along the borders of sanctuary zones) was impacting prey availability for this species (Pichegru et al. 2012). This finding suggests the need for larger sanctuary size in this instance.

Habitat protection and ecosystem resilience In maintaining biodiversity, productivity within a system increases four-fold and enhances the systems recovery potential (Worm et al. 2006). In contrast, when biodiversity in a system declines, stability and water quality decreases exponentially and can lead to resource collapse. For example, sanctuaries in the Mediterranean exhibit higher functional diversity than unprotected areas, emphasizing the importance of sanctuaries at both the species level and the level of trophic group composition (Villamor & Becerro 2012). Analysis of ecosystem-based management approaches reveals large stretches of coastal oceans where sanctuaries play a major role in reducing cumulative impacts and thus improve overall ocean condition (Halpern et al. 2010). For example, spatial planning for gene flow in sanctuaries has been shown to help reduce population isolation and inbreeding and contribute to the resilience of macro algal forests in Tasmania (Durrant et al. 2014). Benefits also extend beyond biotic interactions to include protection from abiotic factors such as increasing sea surface temperatures. In parts of the GBR, that have experienced rising temperatures, sanctuaries still provide conservation outcomes despite these changes (Ban et al. 2012). Sanctuaries also enhance the capacity of coral reefs to withstand flood impacts. In eastern Australia, reefs within sanctuaries resist the impact of perturbation in comparison to fished reefs, and changes on fished reefs are correlated with the magnitude of the flood impact (Olds et al. 2014). Recent research on the GBR found that while marine sanctuaries cannot protect coral reefs directly from acute regional-scale disturbance from a strong tropical cyclone, the impacted reefs within marine sanctuaries supported higher biomass of key fishery-targeted species than fished reefs and should provide valuable sources of larvae to enhance population

recovery and long-term persistence (Emslie et al 2015). Research in the Adriatic Sea showed higher levels of food web complexity, as well as higher biomass for pelagic/demersal and fish/invertebrate ratios in protected areas than exploited systems (Libralato et al. 2010). These results were attributed to the impact of fishing-induced changes in exploited systems. Further, sanctuaries benefit overall ecosystem health and resilience: for example, outbreaks of coral-eating crown-of-thorns starfish are less frequent on no-take reefs, which consequently have higher abundances of coral, the foundation of reef ecosystems (McCook et al. 2010). Additionally, herbivory and coral recruitment are critical ecological processes that underpin reef resilience, and are greater in sanctuary areas and further enhanced on sanctuary reefs near mangroves. Sanctuaries have been shown to provide resistance to the impacts of a changing climate. Sanctuary sites are distinguished from fished sites by showing greater stability in aspects of biodiversity, recovery of large-bodied temperate species, resistance to colonization by sub-tropical vagrants and less pronounced increases in the communityaveraged temperature affinity (Bates et al. 2014). Further, the capacity of sanctuaries to mitigate external disturbances and promote ecological resilience will be critical to resisting an increased frequency of climate-related disturbance (Olds et al. 2014).

Age and size do matter The age of sanctuary zones is important for achieving long-term conservation goals. Older sanctuaries show better results than younger sanctuaries, with densities of fish increasing by approximately 5% per year in sanctuaries compared to unprotected areas (Molloy et al. 2009). Further, densities and age structure of fish are less reliable in young sanctuaries (<10 years) than older sanctuaries (>15 years). In Hawaii, the duration of protection in sanctuaries has significant effects on mean fish length, abundance and fish maturity (Sackett et al. 2014). Additionally, the benefit of protection zones to certain sex-changing fish becomes evident in older sanctuaries (>10 years) and after multiple generations, highlighting the importance of the long-term nature of protection zones (Molloy et al. 2008). Global analysis shows that older sanctuaries are more effective at preventing coral loss, with new sanctuaries exhibiting declines, and these losses stabilizing over time with protection (Selig & Bruno 2010).

Marine protection benefits biodiversity irrespective of sanctuary size (Halpern 2003), however various studies have highlighted an enhanced positive relationship between benefits and sanctuary size. Large sanctuaries are able to accommodate ecosystem interactions without many of the problems associated with networks of smaller sanctuary zones, where

the connectivity between sites can be affected by human activities (Wilhelm et al. 2014). Migratory marine species face a diverse range of pressures during their extensive movements, and are now among the most threatened animals (Lascelles et al. 2014). These wide-ranging species, including many charismatic organisms like marine mammals, seabirds, turtles, sharks, and tuna, also exhibit considerable site fidelity and well-defined habitats, which are prime candidates for enhanced management such as sanctuary designation. Small sanctuaries can be effective in protecting endangered animals that exhibit high levels of site fidelity (Mason & Lowe 2010), such as dusky groupers in the Azores which reside within the small marine sanctuary year-round (Afonso et al. 2011). However, for wide-ranging species such as sharks, sanctuary zones do need to be of the scale to cover their extensive movements (McCook et al. 2010). Protection can also be provided by sanctuary zones that cover important life history stages for species, such as the waved albatross in the Galapagos, that utilize the sanctuaries during nesting, incubation and brooding, with their flight range contracting during these periods (Anderson et al. 2003). Further, analysis of snapper home ranges in New Zealand found residency patterns within sanctuaries that suggest some aspect of the planning results in extreme residency for this species, making sanctuaries effective in protecting species fish stocks (Parsons et al. 2010). The recent creation of large sanctuary zones worldwide has greatly enhanced the chance of achieving global protection targets, as large areas typically contain several ecosystems and habitats that interact ecologically, and allow for more holistic conservation (Wilhelm et al. 2014). In addition, there are recognised opportunities for implementing large sanctuary zones in the pelagic ocean, including weak private property rights, limited habitat transformation and potential lower costs of protected area management (Game et al. 2009). The implementation of large pelagic sanctuaries have been acknowledged as a way of advancing solutions to the broad challenge of conservation planning and management of dynamic systems. In addition, research has shown that the running costs of sanctuaries decrease as the size of the sanctuary increases (Balmford et al. 2004).

Benefits for fisheries Although the major focus of sanctuaries is the protection of biodiversity, there are additional recognized benefits to fisheries. Exploitation of the marine environment by intensive fishing efforts is well documented, and following are just a few examples of this exploitation. Global fish catches have been declining for a number of decades due to a long history of unsustainable fishing practices that have resulted in fisheries collapse and degraded

ecosystems (Pauly et al. 2005). Research reveals that these declines are not confined to the commercial fishing sector. In the USA, analysis of 55 targeted species has revealed that the recreational harvest is increasingly more important than commercial harvest for 71% of the species examined (Ihde et al. 2011). Data from Canada estimates that the recreational fish harvest around the world may represent approximately 12 percent of the global fish harvest (Cooke & Cowx 2004). In Chile, three of the largest reef fish targeted by spearfishers (98% of landings) have been depleted in terms of abundance and size, and catches are moving from large carnivore species to smaller omnivore or herbivore species (Godoy et al. 2010). The benefit of sanctuaries to fisheries can also be revealed when an amnesty on fishing protection is applied. Prior to an amnesty, a sanctuary area in California had 86% higher density of abalone inside the sanctuary zone in comparison to nearby fished areas (Rogers-Bennett et al. 2013). After 3 years of resumed fishing efforts, abalone numbers within the sanctuary were driven down to the point that there was no difference inside or outside the sanctuary zone. In contrast, there are many examples in the literature where protected areas provide benefits to fisheries, in terms of â&#x20AC;&#x153;spillover effectsâ&#x20AC;? and increased larval recruitment.

Spillover effects and larval recruitment Extensive research exists regarding the benefit of sanctuaries in providing spillover effects to surrounding fisheries. Worldwide analysis of spillover effects from 14 no-take sanctuaries suggested local fisheries surrounding 12 of the sanctuaries would be unsustainable without the sanctuaries (Halpern et al. 2009). Further, any negative impacts of the closure were offset by spillover from within the sanctuary. A global synthesis of the literature revealed areas outside reserves did not show declines in biological measures following reserve establishment (Lester et al. 2009). In fact, outside areas either exhibited no change or, in the case of biomass, a significant increase after the reserve was in place. This increase suggests that reserves may benefit outside areas, possibly through the spillover of adults or the export of larvae over the long term.

Sanctuaries in Hawaii exhibit a boundary effect, indicative of spillover effects, whilst also having five times the density of targeted sized yellow tang and 48% higher density of adults than areas open to fishing (Williams et al. 2009a). Modelling based on average increases of biomass within sanctuaries suggests that the benefit to fisheries can accrue up to approximately 50% of total stock area through exported production (Halpern et al. 2004). Recovery of sea urchins in southern China in response to a sanctuary indicates the impact of

removing fishing from this ecosystem in terms of spillover of larvae to surrounding areas (Lau et al. 2011). In the Mediterranean, catch per unit effort of total fish and mean length increased significantly close to the sanctuary for common pandora and red mullet (Stelzenmüller et al. 2007). In South Africa, catch per unit effort increased immediately in the vicinity of a new sanctuary and the increase continued after five years (the time lag for larval recruitment) (Kerwath et al. 2013). This effectively doubled the pre-sanctuary catch per unit effort after 10 years, indicating no detrimental impact of area closure to fishing activities. Fisheries in waters adjacent to sanctuaries in St Lucia increased between 46-90% within five years of creation (Roberts et al. 2001) and world-record sized fish have increased for recreational fishers in Florida since the designation of sanctuary zones (Johnson et al. 1999).

Successful recoveries of lobster fisheries are a great example of the benefits of sanctuary zones to fisheries. In the UK, rapid and large increases in the abundance and sizes of legalsized lobsters within the sanctuary and evidence of spillover of sublegal lobsters to adjacent areas (Sainte-Marie et al. 2011). In the Mediterranean, lobsters in sanctuaries have higher fecundity, mature females are more abundant, egg production is greater and two decades of protection resulted in biomass spillover of an additional 11% of the annual catch and a sixfold increase in egg production (Díaz et al. 2011). Over 8-17 years of protection the harvested spillover of lobsters in the Mediterranean offset the loss of fishing grounds with a net benefit of 10% of the catch in weight (Goñi et al. 2009).

Increased larval recruitment is an important factor in powering the observed spillover effects from sanctuaries. Evidence of larval dispersal in Papua New Guinea has revealed that networks of sanctuaries can function to sustain resident populations both by local replenishment and through larval dispersal from other sanctuaries (Planes et al. 2009). In Mexico, larval retention in sanctuaries results in up to three-fold increases in juvenile recruitment from sanctuaries to fished areas (Cudney-Bueno et al. 2009). Larval export from sanctuaries in the GBR is also significant, with sanctuaries accounting for 28% of the area, but responsible for approximately 50% of juvenile recruitment within the study area (Harrison et al. 2012).

Debate over sanctuaries There remains debate in the literature regarding whether the “spillover effect” is a product of sanctuary designation, or whether the same results can be achieved using traditional fisheries

management approaches (Tupper et al. 2002, Buxton et al. 2014). However, the argument against sanctuaries for fisheries management ignores the important role they play in protecting non-target species and habitats. Sanctuaries and fisheries management tools can produce similar yields for targeted species, however sanctuaries outperform in protecting populations of sedentary species (invertebrates and reef fish) and habitats, making them the preferred method for biodiversity conservation (Hastings & Botsford 1999). Fisheries protection measures are often approached from the perspective of a single economically important species. However, poor stock estimation, improved gear technology and inaccurate reporting by some fishers can result in flawed management plans for targeted species (Sumaila et al. 2000). In addition, even if a fishery is well managed, species that are not subject to management plans still suffer the effects of fishing pressures and are vulnerable as by-catch in other fisheries (Sumaila et al. 2000, Russ & Alcala 2004). Well enforced no-take sanctuaries prevent extractive activities from reducing both the complexity of the habitat and the associated biodiversity (Sumaila et al. 2000).

From a local perspective, Australia prides itself on effective fisheries management (Kearney et al. 2012). However, the truth is that not all fisheries in Australia are well managed. For example, in NSW evidence shows 7.5% of assessed finfish and shark species are over-fished, a number of species are listed as threatened, and 52% of species have their status as undefined or uncertain (Gladstone 2014). In Western Australia, western rock lobster, iconic reef fish such as dhufish and bald chin groper, and most recently Australian herring, have seen such reductions in abundance that fishing mortality was radically reduced (typically by 50%). These reductions were necessary despite â&#x20AC;&#x153;best practiceâ&#x20AC;? fisheries management. Many studies rightly emphasise the need for complimentary management strategies to conserve biodiversity and promote fisheries (Game et al. 2009). By integrating large-scale networks of marine sanctuaries with fishery management, fishery declines can be reversed and urgently needed protection for marine species and their habitats is provided (Gell & Roberts 2003). The creation of networks of marine sanctuaries is viewed as an essential component of marine management (Lubchenco et al. 2003) because it focuses on the protection of the ecosystem rather than managing specific threats or species in isolation (Agardy 2000). Thus, the establishment of no-take sanctuaries should result in improved fisheries in terms of conservation and economic objectives (Yamazaki et al. 2012). Recent advances across conservation, oceanography and fisheries science provide the evidence, tools and information

to confirm sanctuaries as defensible and feasible instruments for conservation (Game et al. 2009).

Socio-economic considerations This review highlights some of the socio-economic benefits of sanctuaries, such as tourism, research and management but is by no a means a comprehensive review of literature pertaining to these areas of research. Tourism income Globally, there are many examples of sanctuaries resulting in economic gains for the tourism industry. For example, investigation of the live-aboard dive industry in the Great Barrier Reef and Coral Sea reported three key finding: (1) Each year live-aboard dive boats are directly responsible for generating at least AU$16 million worth of income in the Cairns/Port Douglas region; (2) Visitors participating in different types of trips gain their highest levels of 'satisfaction' from interacting with different types of species; and (3) Visitors to Osprey Reef would be willing to pay more for a 'guaranteed' sighting of sharks than they would for a 'guaranteed' sighting of large fish, marine turtles or a 'wide variety of species' (Stoeckl et al. 2010). Given the previously described benefits to biodiversity and shark species that sanctuaries provide, these economic and tourism benefits relate directly to the provision of sanctuary zones. Sanctuaries in NSW provide another example of tourism income, where juvenile and sub-adult grey nurse sharks have re-established at former aggregation sites since the areas became sanctuaries. Results show sanctuaries are assisting the recovery of the east coast population, as well as providing tourism income to local dive operators whose dive trips have increased at these sites four-fold since the return of the grey nurse sharks (Lynch et al. 2013). NSW provides more evidence of the benefits to tourism of sanctuary zones, with the establishment of the Solitary Island Marine Park seeing a 20% increase in local businessâ&#x20AC;&#x2122;s turnover in the first five years. Jervis Bay Marine Park has brought an estimated $2.4 million into the region through marine tourism (Hoisington 2013). Abroad, shark diving tourism in Palau generates US$18 million per year and 8% of the GDP (Vianna et al. 2012). Whereas if the population of approximately 100 sharks that interact with divers were to be harvested by fishers their value would be approximately US$10,800. Thus, protection of the shark populations provides an economic benefit to multiple stakeholders and ensures their ecological sustainability. Finally, cost-benefit analysis of protecting the environment and

tourism values of the GBR suggest that relative to the revenue generated by reef tourism, current expenditure on protecting the reef is minor (McCook et al. 2010).

A review of the social impacts of marine reserves in New Zealand found that while there can be an initial negative impact on commercial and recreational fishing, over time there are benefits that relate primarily to increases in visitor numbers and recreation activity which have flow-on benefits for the local community as well as demands for infrastructure such as roads, parking and waste facilities in the vicinity of the reserve and tourism businesses (Taylor and Buckenham, 2003). They found that opposition to marine reserves comes largely from commercial and recreational fishers who perceive that they will have their activities adversely affected by the establishment of a reserve. However, the views of a majority of fishers can change in favour of reserves with time (Taylor and Buckenham, 2003).

Benefit to science Sanctuaries provide important baseline sites to compare with fished areas, due to both the direct and indirect changes that occur when fishing is prevented. In other words, they provide the control to the experiment of human activities on marine ecosystems (Ballantine 2014). They show the effects of fishing on ecosystems through depleted populations, habitat change and trophic cascades (Costello 2014). For example, New Zealand has some of the oldest and best-studied sanctuaries, which provide comparable replicates for studying the recovery of previously exploited habitats, and shows the re-establishment of trophic controls over community structure and productivity (Langlois & Ballantine 2005). Older sanctuaries can be used to test scientific questions, as the protection afforded by the marine sanctuary allows investigation into the depletion from fishing pressure and how this changes upper-level predators and trophic processes between habitats. This emphasizes the importance of longevity when creating sanctuaries, as changes to ecosystems are likely continuous and will vary with environments, climate and species distributions (Leleu et al. 2012). Research in the Pacific has highlighted the importance of replicating no-take sanctuaries to reduce the risk of overexploitation by fisheries and to provide scientific testing of fishery and conservation management strategies (Murray et al. 1999).

Economic management Sanctuary management costs are substantially lower relative to fisheries subsidies. From the 53 countries that contribute 95% of global fisheries catch, sanctuary subsidies to fisheries

total US$870 million, just 2.5-2.8% of the total fisheries subsidies (US$30-34 billion) paid annually for the maintenance of the biodiversity that sustains them (Cullis-Suzuki & Pauly 2010). On a local scale, analysis of the impact of re-zoning in Australiaâ&#x20AC;&#x2122;s Moreton Bay suggests that, counter to initial claims, non-market recreational fishing benefits may have increased by between $1.3 million and $2.5 million a year, with a current total annual value of around $20 million (Pascoe et al. 2014). A broad-scale study based on the running costs of 83 sanctuaries worldwide was conducted to determine the estimated costs of a global network of marine protected areas (Balmford et al. 2004). Models estimated that a global sanctuary network that meets the World Parks Congress target of conserving 20â&#x20AC;&#x201C;30% of the world's seas would cost between $5 billion and $19 billion annually to run and would probably create around one million jobs. These gross network costs, although substantial, are actually less than current government expenditures on industrial fisheries subsidies.

Community Support for sanctuaries In general, the wider public endorses marine sanctuaries as important tools for biodiversity conservation. A global survey of 7000 people, conducted by the Zoological Society of London, revealed that on average people want 50% of global oceans set aside for conservation protection compared to the current level of 3% protection (ZSL 2014). A survey of the New Zealand public, showed that they believed 30% of the marine environment is currently protected and that 36% should be protected with sanctuaries, but in fact 0.3% is protected, indicating public would support further marine protection (Eddy 2014). Research into stakeholder perceptions of a sanctuary in the UK indicated that the social, economic and environmental benefits of the sanctuaries were perceived to outweigh any costs (Rees et al. 2013).

A global review of commercial fishers reactions to sanctuary implementation for fishery management purposes showed general acceptance of sanctuaries as effective and beneficial management tools (Pita et al. 2011). In the GBR, approximately 56% of fishers reported losing at least one of their regular fishing locations under the new zoning plan in 2004. However, fishers generally compensated by shifting their fishing effort to other areas they knew to be good fishing locations, and by finding new areas that they had not exploited previously (De Freitas et al. 2013). Establishment of sanctuaries in Hawaii showed that fishing effort was displaced, however potential economic losses were offset by expanded operating ranges and favourable market forces, which meant fisher socioeconomic well being

was not compromised (Stevenson et al. 2013). In New Zealand, the majority of respondents supported the sanctuaries after public involvement in the planning process and appropriate communication to help reduce conflicts (Wolfenden et al. 1994). Only 25% of fisheries objectives and 30% of biodiversity objective were considered to be sources of conflict in sanctuary designs, suggesting that better segregation of activities in planning could resolve most potential conflicts (Rice et al. 2012). Management restrictions in Madagascar were more readily supported by fishers with experience in decision-making, access to livelihood alternatives and more permanent housing, and they were also more likely to be opinionated (McClanahan et al. 2014).

Recreational fishers from the GBR were surveyed three years after the new zoning came in and 68% believed a change to zoning was a good idea, with 57% endorsing the actual new zones (Sutton & Tobin 2009). The majority believed in the conservation values of the new zones and did not feel it would affect their recreational fishing activity. For repeat visitors to Ningaloo Marine Park, 87% could correctly identify the location of the nearest sanctuary zone; for first time visitors this dropped to 52% (Smallwood & Beckley 2012). About 75% of both visitor types stated that these zones had not affected their fishing activity. Sanctuaries in West Hawaii were moderately effective at alleviating conflict between different stakeholders, where dive operators view commercial fishers and aquarium-trade fisherman as a threat to the coral reef ecosystem (Stevenson & Tissot 2013). In Puerto Rico, 80% of artisanal fisherman, 73% of concessionaires (ie. SCUBA diving, charter boats etc.) and 52% of registered vessel owners supported sanctuaries, recognizing declines in coral reef condition and water quality as the main drivers for protection (Hernรกndez-Delgado et al. 2014).

Compliance Levels of compliance in sanctuary zones are a good way of assessing fishers support for this conservation measure. A survey of recreational fishers in the GBR indicated that the majority (90%) comply with sanctuary zones (Arias & Sutton 2013). Likewise, most (92%) reported not personally knowing anyone who had intentionally fished in a sanctuary zone, indicating that fishers' perceive high levels of compliance among their peers. They were motivated to comply with no-take sanctuaries primarily by their beliefs about penalties for noncompliance, followed by beliefs about the fishery benefits of sanctuaries. Conversely, despite support for biodiversity conservation measures and fisheries management regulations being high, non-compliance with minimum size limits was common in south western Australia and

further education of anglers appears to be necessary (Prior & Beckley 2007). In Tanzania, positive signs for communities from local conservation efforts of sanctuaries include increased catch/income, awareness and compliance (Yakupitiyage & Machumu 2013). Fishers in Mexico were able to identify the key objectives of sanctuaries for regulation, conservation and economic improvement. Further, they cared about ecosystem sustainability and endorsed the initiatives while expecting additional benefits. They were concerned about illegal fishing once the sanctuary zones came into effect and were happy to be part of enforcing the sanctuaries (Velez et al. 2014).

CONCLUSION There is extensive support in the scientific literature for the benefits of marine sanctuaries. Marine sanctuaries are effective tools for providing protection for biodiversity at both species and ecosystem scales. Their implementation increases ecosystem resilience to threatening processes and has been recognized as complementary to other management strategies for preserving the marine environment. Networks of well-connected, large marine sanctuaries should be the preferred and long-term objective for achieving conservation outcomes and meeting environmental targets for protecting biodiversity. While sanctuaries cannot provide direct protection from threatening processes such as climate change, water pollution and invasive species, their ability to promote and protect functional diversity means these ecosystems are more likely to withstand these other threats. In addition to conservation outcomes, sanctuaries provide important contributions to fisheries and the economy, and are well supported by broad range of stakeholders. Marine sanctuaries should be used in conjunction with fisheries management plans, in order to protect target species, non-target species and their habitats. Both approaches are necessary to ensure the sustainable future of our oceans from both a conservation and an economic perspective. Given the 2014 target set by the World Parks Congress to protect 30% of the global marine environment with sanctuaries by 2030, and given the multitude of benefits associated with these sanctuaries, retaining the current National Representative System of Marine Protected Areas in Australia would be a defensible and wise decision.

REFERENCES Aburto-Oropeza O, Erisman B, Galland GR, Mascareñas-Osorio I, Sala E, Ezcurra E (2011) Large recovery of fish biomass in a no-take marine reserve. PLoS One 6:e23601 Afonso P, Fontes J, Santos RS (2011) Small marine reserves can offer long term protection to an endangered fish. Biol Conserv 144:2739-2744 Agardy T (2000) Information needs for marine protected areas: scientific and societal. Bulletin of Marine Science 66:875-888 Alves MDdO, Schwamborn R, Borges JCG, Marmontel M, Costa AF, Schettini CAF, Araújo MEd (2013) Aerial survey of manatees, dolphins and sea turtles off northeastern Brazil: Correlations with coastal features and human activities. Biol Conserv 161:91100 Anderson DJ, Huyvaert KP, Wood DR, Gillikin CL, Frost BJ, Mouritsen H (2003) At-sea distribution of waved albatrosses and the Galápagos Marine Reserve. Biol Conserv 110:367-373 Arias A, Sutton SG (2013) Understanding recreational fishers' compliance with no-take zones in the Great Barrier Reef Marine Park. Ecology and Society 18:1-9 Ault JS, Smith SG, Bohnsack JA, Luo J, Zurcher N, McClellan DB, Ziegler TA, Hallac DE, Patterson M, Feeley MW, Ruttenberg BI, Hunt J, Kimball D, Causey B (2013) Assessing coral reef fish population and community changes in response to marine reserves in the Dry Tortugas, Florida, USA. Fisheries Research 144:28-37 Ballantine B (2014) Fifty years on: Lessons from marine reserves in New Zealand and principles for a worldwide network. Biol Conserv Balmford A, Gravestock P, Hockley N, McClean CJ, 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 Ban NC, Natalie C BAN, Robert L P, Scarla W (2012) Conservation Objectives and SeaSurface Temperature Anomalies in the Great Barrier Reef; Objetivos de Conservación y Anomalías en la Temperatura de la Superficie del Mar en la Gran Barrera Arrecifal. Conservation Biology 26:799-809 Bates AE, Bates AE, Barrett NS, Stuart-Smith RD, Holbrook NJ (2014) Resilience and signatures of tropicalization in protected reef fish communities. Nature Climate Change 4:62-67 Berkeley SA, Hixon MA, Larson RJ, Love MS (2004) Fisheries sustainability via protection of age structure and spatial distribution of fish populations. Fisheries 29:23-32

Branch GM, Odendaal F (2003) The effects of marine protected areas on the population dynamics of a South African limpet, Cymbula oculus, relative to the influence of wave action. Biol Conserv 114:255-269 Butcher PA, Boulton AJ, Macbeth WG, Malcolm HA (2014) Long-term effects of marine park zoning on giant mud crab Scylla serrata populations in three Australian estuaries. Marine Ecology Progress Series 508:163-176 Buxton CD, Hartmann K, Kearney R, Gardner C (2014) When is spillover from marine reserves likely to benefit fisheries? PloS one 9:e107032 Cooke SJ, Cowx IG (2004) The Role of Recreational Fishing in Global Fish Crises. BioScience 54:857-859 Costello MJ (2014) Long live Marine Reserves: A review of experiences and benefits. Biol Conserv 176:289-296 Cudney-Bueno R, Lavín MF, Marinone SG, Raimondi PT, Shaw WW (2009) Rapid effects of marine reserves via larval dispersal. PloS One 4:e4140 Cullis-Suzuki S, Pauly D (2010) Marine protected area costs as “beneficial” fisheries subsidies: a global evaluation. Coastal Management 38:113-121 De Freitas DM, Sutton SG, Moloney JM, Lédée EJ, Tobin RC (2013) Spatial substitution strategies of recreational fishers in response to zoning changes in the Great Barrier Reef Marine Park. Marine Policy 40:145-153 Díaz D, Mallol S, Parma AM, Goñi R (2011) Decadal trend in lobster reproductive output from a temperate marine protected area. Marine Ecology Progress Series 433:149-157 Dudley N (2008) Guidelines for applying protected area management categories. IUCN Durrant H, Burridge CP, Kelaher BP, Barrett NS, Edgar GJ, Coleman MA (2014) Implications of macroalgal isolation by distance for networks of marine protected areas. Conservation Biology 28:438-445 Eddy TD (2014) One hundred-fold difference between perceived and actual levels of marine protection in New Zealand. Marine Policy 46:61-67 Edgar GJ, Stuart-Smith RD, Willis TJ, Kininmonth S, Baker SC, Banks S, Barrett NS, Becerro MA, Bernard AT, Berkhout J (2014) Global conservation outcomes depend on marine protected areas with five key features. Nature Emslie MJ, Logan M, Williamson DH, Ayling T, MacNeil MA, Ceccarelli D, Cheal AJ, Evans RD, Johns KA, Jonker MJ, Miller IR, Osborne K, Russ GR, Sweatman HPA (2015) Expectations and outcomes of reserve network performance following rezoning of the Great Barrier Reef Marine Park. Current Biology. In Press

Florin AB, Bergström U, Ustups D, Lundström K, Jonsson PR (2013) Effects of a large northern European no-take zone on flatfish populationsa. Journal of Fish Biology 83:939-962 Follesa MC, Cannas R, Cau A, Cuccu D, Gastoni A, Ortu A, Pedoni C, Porcu C, Cau A (2011) Spillover effects of a Mediterranean marine protected area on the European spiny lobster Palinurus elephas (Fabricius, 1787) resource. Aquatic Conservation: Marine and Freshwater Ecosystems 21:564-572 Game ET, Grantham HS, Hobday AJ, Pressey RL, Lombard AT, Beckley LE, Gjerde K, Bustamante R, Possingham HP, Richardson AJ (2009) Pelagic protected areas: the missing dimension in ocean conservation. Trends in Ecology & Evolution 24:360-369 Gell FR, Roberts CM (2003) Benefits beyond boundaries: the fishery effects of marine reserves. Trends in Ecology & Evolution 18:448-455 Gladstone W (2014) Criticisms of science, social impacts, opinion leaders, and targets for notake zones led to cuts in New South Wales. Aquatic Conservation 24:287 Godoy N, Gelcich S, Vásquez JA, Castilla JC (2010) Spearfishing to depletion: evidence from temperate reef fishes in Chile. Ecological Applications 20:1504-1511 Goetze JS, Fullwood LAF (2013) Fiji’s largest marine reserve benefits reef sharks. Coral Reefs 32:121-125 Goñi R, Hilborn R, Díaz D, Mallol S, Adlerstein S (2009) Net contribution of spillover from a marine reserve to fishery catches. Marine Ecology Progress Series 400:233 Halpern BS (2003) The impact of marine reserves: do reserves work and does reserve size matter? Ecological Applications 13:117-137 Halpern BS, Gaines SD, Warner RR (2004) Confounding effects of the export of production and the displacement of fishing effort from marine reserves. Ecological Applications 14:1248-1256 Halpern BS, Lester SE, Kellner JB (2009) Spillover from marine reserves and the replenishment of fished stocks. Environmental Conservation 36:268-276 Halpern BS, Lester SE, McLeod KL (2010) Placing marine protected areas onto the ecosystem-based management seascape. Proceedings of the National Academy of Sciences 107:18312-18317 Harasti D, Martin-Smith K, Gladstone W (2014) Does a no-take marine protected area benefit seahorses? PloS One 9:e105462 Harrison HB, Williamson DH, Evans RD, Almany GR, Thorrold SR, Russ GR, Feldheim KA, Van Herwerden L, Planes S, Srinivasan M (2012) Larval export from marine

reserves and the recruitment benefit for fish and fisheries. Current Biology 22:10231028 Hastings A, Botsford LW (1999) Equivalence in yield from marine reserves and traditional fisheries management. Science 284:1537-1538 Hernández-Delgado EA, Shivlani M, Sabat AM (2014) Ecosystem-Based and CommunityBased Model Integration to Designate Coral Reef No-Take Marine Protected Areas: A Case Study from Puerto Rico. Natural Resources 5:538-560 Hoisington C (2013) The Marine Protection Dividend: NSW marine parks deliver more over time. Centre for Policy Development Occasional Paper:4 Hooker SK, Gerber LR (2004) Marine reserves as a tool for ecosystem-based management: the potential importance of megafauna. BioScience 54:27-39 Ihde TF, Wilberg MJ, Loewensteiner DA, Secor DH, Miller TJ (2011) The increasing importance of marine recreational fishing in the US: Challenges for management. Fisheries Research 108:268-276 Jensen OP, Ortega-Garcia S, Martell SJ, Ahrens RN, Domeier ML, Walters CJ, Kitchell JF (2010) Local management of a “highly migratory species”: the effects of long-line closures and recreational catch-and-release for Baja California striped marlin fisheries. Progress in Oceanography 86:176-186 Johnson DR, Funicelli NA, Bohnsack JA (1999) Effectiveness of an Existing Estuarine NoTake Fish Sanctuary within the Kennedy Space Center, Florida. North American Journal of Fisheries Management 19:436-453 Kearney R, Buxton CD, Farebrother G (2012) Australia’s no-take marine protected areas: Appropriate conservation or inappropriate management of fishing? Marine Policy 36:1064-1071 Kelly S, Scott D, MacDiarmid AB, Babcock RC (2000) Spiny lobster, Jasus edwardsii, recovery in New Zealand marine reserves. Biol Conserv 92:359-369 Kerwath SE, Winker H, Götz A, Attwood CG (2013) Marine protected area improves yield without disadvantaging fishers. Nature communications 4:2347 Knip DM, Heupel MR, Simpfendorfer CA (2012) Evaluating marine protected areas for the conservation of tropical coastal sharks. Biol Conserv 148:200-209 Langlois TJ, Ballantine WJ (2005) Marine Ecological Research in New Zealand: Developing Predictive Models through the Study of No‐Take Marine Reserves. Conservation Biology 19:1763-1770

Lascelles B, Notarbartolo Di Sciara G, Agardy T, Cuttelod A, Eckert S, Glowka L, Hoyt E, Llewellyn F, Louzao M, Ridoux V, Tetley MJ (2014) Migratory marine species: their status, threats and conservation management needs. Aquatic Conservation: Marine and Freshwater Ecosystems 24:111-127 Lau DC, Dumont CP, Lui G, Qiu J-W (2011) Effectiveness of a small marine reserve in southern China in protecting the harvested sea urchin< i> Anthocidaris crassispina</i>: A mark-and-recapture study. Biol Conserv 144:2674-2683 Leleu K, Remy-Zephir B, Grace R, Costello MJ (2012) Mapping habitats in a marine reserve showed how a 30-year trophic cascade altered ecosystem structure. Biol Conserv 155:193-201 Lester SE, Halpern BS, Grorud-Colvert K, Lubchenco J, Ruttenberg BI, Gaines SD, AiramĂŠ S, Warner RR (2009) Biological effects within no-take marine reserves: a global synthesis. Marine Ecology Progress Series 384:33-46 Libralato S, Coll M, Tempesta M, Santojanni A, Spoto M, Palomera I, Arneri E, Solidoro C (2010) Food-web traits of protected and exploited areas of the Adriatic Sea. Biol Conserv 143:2182-2194 Ling S, Johnson C (2012) Marine reserves reduce risk of climate-driven phase shift by reinstating size-and habitat-specific trophic interactions. Ecological Applications 22:1232-1245 Ling S, Johnson C, Frusher S, Ridgway K (2009) Overfishing reduces resilience of kelp beds to climate-driven catastrophic phase shift. Proceedings of the National Academy of Sciences 106:22341-22345 Lubchenco J, Palumbi SR, Gaines SD, Andelman S (2003) Plugging a hole in the ocean: the emerging science of marine reserves 1. Ecological applications 13:3-7 Ludynia K, Kemper J, Roux J-P (2012) The Namibian Islandsâ&#x20AC;&#x2122; Marine Protected Area: Using seabird tracking data to define boundaries and assess their adequacy. Biol Conserv 156:136-145 Lynch TP, Harcourt R, Edgar G, Barrett N (2013) Conservation of the Critically Endangered Eastern Australian Population of the Grey Nurse Shark (Carcharias taurus) Through Cross-Jurisdictional Management of a Network of Marine-Protected Areas. Environmental management 52:1341-1354 Maggs JQ, Mann BQ, Cowley PD (2013) Contribution of a large no-take zone to the management of vulnerable reef fishes in the South-West Indian Ocean. Fisheries Research 144:38-47

Mason TJ, Lowe CG (2010) Home range, habitat use, and site fidelity of barred sand bass within a southern California marine protected area. Fisheries Research 106:93-101 McClanahan TR, Cinner JE, Abunge C, Rabearisoa A, Mahatante P, Ramahatratra F, Andrianarivelo N (2014) Perceived Benefits of Fisheries Management Restrictions in Madagascar. Ecology and Society 19:5 McCook LJ, Ayling T, Cappo M, Choat JH, Evans RD, De Freitas DM, Heupel M, Hughes TP, Jones GP, Mapstone B (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 107:18278-18285 Molloy PP, McLean IB, Côté IM (2009) Effects of marine reserve age on fish populations: a global meta-analysis. Journal of Applied Ecology 46:743-751 Molloy PP, Reynolds JD, Gage MJG, Mosqueira I, Côté IM (2008) Links between sex change and fish densities in marine protected areas. Biol Conserv 141:187-197 Murray SN, Ambrose RF, Bohnsack JA, Botsford LW, Carr MH, Davis GE, Dayton PK, Gotshall D, Gunderson DR, Hixon MA (1999) No-take reserve networks: sustaining fishery populations and marine ecosystems. Fisheries 24:11-25 Olds AD, Pitt KA, Maxwell PS, Babcock RC, Rissik D, Connolly RM (2014) Marine reserves help coastal ecosystems cope with extreme weather. Global Change Biology 20:3050-3058 Parsons DM, Morrison MA, Slater MJ (2010) Responses to marine reserves: Decreased dispersion of the sparid Pagrus auratus (snapper). Biol Conserv 143:2039-2048 Pascoe S, Doshi A, Dell Q, Tonks M, Kenyon R (2014) Economic value of recreational fishing in Moreton Bay and the potential impact of the marine park rezoning. Tourism Management 41:53-63 Pauly D, Watson R, Alder J (2005) Global trends in world fisheries: impacts on marine ecosystems and food security. Philosophical Transactions of the Royal Society B: Biological Sciences 360:5-12 Péron C, Grémillet D, Prudor A, Pettex E, Saraux C, Soriano-Redondo A, Authier M, Fort J (2013) Importance of coastal Marine Protected Areas for the conservation of pelagic seabirds: The case of Vulnerable yelkouan shearwaters in the Mediterranean Sea. Biol Conserv 168:210-221 Pichegru L, Grémillet D, Crawford R, Ryan P (2010) Marine no-take zone rapidly benefits endangered penguin. Biology Letters 6:498-501

Pichegru L, Ryan PG, Van Eeden R, Reid T (2012) Industrial fishing, no-take zones and endangered penguins. Biol Conserv 156:117 Pillans S, Ortiz J-C, Pillans RD, Possingham HP (2007) The impact of marine reserves on nekton diversity and community composition in subtropical eastern Australia. Biol Conserv 136:455-469 Pita C, Pierce G, Theodossiou I, Macpherson K (2011) An overview of commercial fishers’ attitudes towards marine protected areas. Hydrobiologia 670:289-306 Planes S, Jones GP, Thorrold SR (2009) Larval dispersal connects fish populations in a network of marine protected areas. Proceedings of the National Academy of Sciences 106:5693-5697 Prior S, Beckley L (2007) Characteristics of recreational anglers in the Blackwood Estuary, a popular tourist destination in Southwestern Australia. Tourism in Marine Environments 4:15-28 Rees SE, Attrill MJ, Austen MC, Mangi SC, Rodwell LD (2013) A thematic cost-benefit analysis of a marine protected area. Journal of Environmental Management 114:476485 Rice J, Moksness E, Attwood C, Brown SK, Dahle G, Gjerde KM, Grefsrud ES, Kenchington R, Ring Kleiven A, McConney P, Ngoile MAK, Næsje TF, Olsen E, Olsen EM, Sanders J, Sharma C, Vestergaard O, Westlund L (2012) The role of MPAs in reconciling fisheries management with conservation of biological diversity. Ocean & Coastal Management 69:217-230 Robbins WD, Hisano M, Connolly SR, Choat JH (2005) Ongoing Collapse of Coral-Reef Shark Populations. Current Biology 16:2314-2319 Roberts CM, Bohnsack JA, Gell F, Hawkins JP, Goodridge R (2001) Effects of marine reserves on adjacent fisheries. Science 294:1920-1923 Rogers-Bennett L, Hubbard KE, Juhasz CI (2013) Dramatic declines in red abalone populations after opening a “de facto” marine reserve to fishing: Testing temporal reserves. Biol Conserv 157:423-431 Ruppert JLW, Travers MJ, Smith LL, Fortin M-J (2013) Caught in the middle: combined impacts of shark removal and coral loss on the fish communities of coral reefs. PloS one 8:e74648 Russ GR, Alcala AC (2004) Marine reserves: long-term protection is required for full recovery of predatory fish populations. Oecologia 138:622-627

Sackett D, Drazen J, Moriwake V, Kelley C, Schumacher B, Misa WXE (2014) Marine protected areas for deepwater fish populations: an evaluation of their effects in Hawai’i. Marine Biology 161:411-425 Sainte-Marie B, Hoskin M, Coleman R, Von Carlshausen E, Davis C (2011) Variable population responses by large decapod crustaceans to the establishment of a temperate marine no-take zone. Canadian Journal of Fisheries and Aquatic Sciences 68:185-200 Schofield G, Scott R, Dimadi A, Fossette S, Katselidis KA, Koutsoubas D, Lilley MKS, Pantis JD, Karagouni AD, Hays GC (2013) Evidence-based marine protected area planning for a highly mobile endangered marine vertebrate. Biol Conserv 161:101109 Selig ER, Bruno JF (2010) A global analysis of the effectiveness of marine protected areas in preventing coral loss. PLoS One 5:e9278 Smallwood CB, Beckley LE (2012) Spatial distribution and zoning compliance of recreational fishing in Ningaloo Marine Park, north-western Australia. Fisheries Research 125–126:40-50 Stelzenmüller V, Maynou F, Martín P (2007) Spatial assessment of benefits of a coastal Mediterranean Marine Protected Area. Biol Conserv 136:571-583 Stevenson TC, Tissot BN (2013) Evaluating marine protected areas for managing marine resource conflict in Hawaii. Marine Policy 39:215-223 Stevenson TC, Tissot BN, Walsh WJ (2013) Socioeconomic consequences of fishing displacement from marine protected areas in Hawaii. Biol Conserv 160:50-58 Stoeckl N, Birtles A, Farr M, Mangott A, Curnock M, Valentine P (2010) Live-aboard dive boats in the Great Barrier Reef: regional economic impact and the relative values of their target marine species. Tourism Economics 16:995-1018 Sumaila UR, Guénette S, Alder J, Chuenpagdee R (2000) Addressing ecosystem effects of fishing using marine protected areas. ICES Journal of Marine Science: Journal du Conseil 57:752-760 Sutton SG, Tobin RC (2009) Recreational fishers' attitudes towards the 2004 rezoning of the Great Barrier Reef Marine Park. Environmental Conservation 36:245-252 Taylor N, Buckenham B (2003) Social Impacts of Marine Reserves in New Zealand. Department of Conservation Trebilco R, Gales R, Baker GB, Terauds A, Sumner MD (2008) At sea movement of Macquarie Island giant petrels: Relationships with marine protected areas and Regional Fisheries Management Organisations. Biol Conserv 141:2942-2958

Tupper MH, Wickstrom K, Hilborn R, Roberts CM, Bohnsack JA, Gell F, Hawkins JP, Goodridge R (2002) Marine Reserves and Fisheries Management. Science 295:12331235 Velez M, Adlerstein S, Wondolleck J (2014) Fishers' perceptions, facilitating factors and challenges of community-based no-take zones in the Sian Ka’an Biosphere Reserve, Quintana Roo, Mexico. Marine Policy 45:171-181 Vianna GMS, Meekan MG, Pannell DJ, Marsh SP, Meeuwig JJ (2012) Socio-economic value and community benefits from shark-diving tourism in Palau: A sustainable use of reef shark populations. Biol Conserv 145:267-277 Villamor A, Becerro MA (2012) Species, trophic, and functional diversity in marine protected and non-protected areas. Journal of Sea Research 73:109-116 Watson JEM, Dudley N, Segan DB, Hockings M (2014) The performance and potential of protected areas. Nature 515:67-73 Wilhelm TA, Sheppard CRC, Sheppard ALS, Gaymer CF, Parks J, Wagner D, Lewis Na (2014) Large marine protected areas – advantages and challenges of going big. Aquatic Conservation: Marine and Freshwater Ecosystems 24:24-30 Williams ID, Walsh WJ, Claisse JT, Tissot BN, Stamoulis KA (2009a) Impacts of a Hawaiian marine protected area network on the abundance and fishery sustainability of the yellow tang, Zebrasoma flavescens. Biol Conserv 142:1066-1073 Williams R, Lusseau D, Hammond PS (2009b) The role of social aggregations and protected areas in killer whale conservation: the mixed blessing of critical habitat. Biol Conserv 142:709-719 Wolfenden J, Cram F, Kirkwood B (1994) Marine reserves in New Zealand: A survey of community reactions. Ocean & Coastal Management 25:31-51 Worm B, Barbier EB, Beaumont N, Duffy JE, Folke C, Halpern BS, Jackson JB, Lotze HK, Micheli F, Palumbi SR (2006) Impacts of biodiversity loss on ocean ecosystem services. Science 314:787-790 Yakupitiyage A, Machumu E (2013) Effectiveness of Marine Protected Areas in Managing the Drivers of Ecosystem Change: A Case of Mnazi Bay Marine Park, Tanzania. Yamazaki S, Grafton QR, Kompas T, Jennings S (2012) Biomass management targets and the conservation and economic benefits of marine reserves. Fish and Fisheries ZSL (2014) Planet’s protected areas fall short of public’s expectations. http://www.zsl.org/conservation/news/planet’s-protected-areas-fall-short-of-public’sexpectations. Zoological Society of London.