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	<front>
		<journal-meta>
			<journal-id journal-id-type="publisher-id">SCIMAR</journal-id>
			<journal-title-group>
				<journal-title>Scientia Marina</journal-title>
				<abbrev-journal-title abbrev-type="publisher">Sci. Mar.</abbrev-journal-title>
			</journal-title-group>
			<issn publication-format="print">0214-8358</issn>
			<issn publication-format="electronic">1886-8134</issn>
			<publisher>
				<publisher-name>Consejo Superior de Investigaciones Cient&#xed;ficas</publisher-name>
			</publisher>
		</journal-meta>
		<article-meta>
			<article-id pub-id-type="publisher-id">scimar.05417.080</article-id>
			<article-id pub-id-type="doi">10.3989/scimar.05417.080</article-id>
			<article-categories>
				<subj-group subj-group-type="heading">
					<subject>News and comments</subject>
				</subj-group>
			</article-categories>
			<title-group>
				<article-title>Evidence of economic benefits from marine protected areas</article-title>
				<trans-title-group xml:lang="es">
					<trans-title>Evidencia de beneficios econ&#xf3;micos de las &#xc1;reas Marinas Protegidas</trans-title>
				</trans-title-group>
			</title-group>
			<contrib-group>
				<contrib contrib-type="author" corresp="yes">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-2362-0328</contrib-id>
					<name>
						<surname>Costello</surname>
						<given-names>Mark John</given-names>
					</name>
					<email xlink:href="mark.j.costello@nord.no">mark.j.costello@nord.no</email>
					<aff id="aff1a"><institution content-type="faculty">Faculty of Biosciences and Aquaculture</institution>, <institution content-type="university">Nord Universitet</institution>, <addr-line>Postboks 1490, 8049 Bod&#xf8;</addr-line>, <country>Norway</country>.</aff>
					<aff id="aff1b"><institution content-type="college">College of Life Science</institution>, <institution content-type="university">Ocean University of China</institution>, <addr-line>Qingdao</addr-line>, <country>China</country>.</aff>
				</contrib>
				<contrib contrib-type="editor">
					<name>
						<surname>Peters</surname>
						<given-names>F.</given-names>
					</name>
				</contrib>
			</contrib-group>
			<pub-date pub-type="epub">
				<day>30</day>
				<month>03</month>
				<year>2024</year>
			</pub-date>
			<pub-date pub-type="collection">
				<month>03</month>
				<year>2024</year>
			</pub-date>
			<volume>88</volume>
			<issue>1</issue>
			<elocation-id>e080</elocation-id>
			<pub-history>
				<event>
					<event-desc>Received</event-desc>				
					<date date-type="received">
						<day>02</day>
						<month>10</month>
						<year>2023</year>
					</date>
				</event>
				<event>
					<event-desc>Accepted</event-desc>				
					<date date-type="accepted">
						<day>24</day>
						<month>10</month>
						<year>2023</year>
					</date>
				</event>
				<event>
					<event-desc>Published</event-desc>				
					<date date-type="pub">
						<day>27</day>
						<month>03</month>
						<year>2024</year>
					</date>
				</event>
			</pub-history>
			<permissions>
				<copyright-statement>&#xa9; 2024 CSIC</copyright-statement>
				<copyright-year>2024</copyright-year>
				<license license-type="open-access" xlink:href="https://creativecommons.org/licenses/by/4.0/">
					<license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution 4.0 International (CC BY 4.0) License.</license-p>
				</license>
			</permissions>
			<self-uri xlink:href="http://scientiamarina.revistas.csic.es/index.php/scientiamarina/article/view/XXXX/XXXX"/>
			<abstract>
				<title>Summary</title>
				<p>Marine protected areas (MPAs) have been used for biodiversity conservation for decades. However, critics argue that evidence of their economic benefits is weak, particularly with regard to fisheries. This continued opposition to MPAs for fisheries slows progress towards conservation targets and undermines the economic and ecological sustainability of the oceans. This paper provides 48 examples of fishery-related and 31 of tourism-related economic benefits in 25 and 24 countries, respectively. There was no evidence of net costs of MPAs to fisheries anywhere. Fishery benefits included increased fish stocks, catch volumes, catch per unit effort, fecundity and larval export, and larger fish and lobsters. Well-designed and enforced MPAs provide sustainable benefits for fishing communities and even sub-optimally designed MPAs can provide economic advantages. MPAs represent one of the best strategies for maintaining the sustainable exploitation of marine resources.</p>
			</abstract>
			<trans-abstract xml:lang="es">
				<title>Resumen</title>
				<p>Las &#xc1;reas Marinas Protegidas (AMP) se han utilizado durante d&#xe9;cadas para la conservaci&#xf3;n de la biodiversidad. Sin embargo, los cr&#xed;ticos argumentan que la evidencia de sus beneficios econ&#xf3;micos es d&#xe9;bil, particularmente en lo que respecta a la pesca. Esta continua oposici&#xf3;n a las AMP en pesquer&#xed;as obstaculiza el progreso hacia los objetivos de las AMP y socava la sostenibilidad econ&#xf3;mica y ecol&#xf3;gica de los oc&#xe9;anos. Este documento proporciona 48 ejemplos de pesquer&#xed;a y 31 de beneficios econ&#xf3;micos relacionados con el turismo en 24 y 23 pa&#xed;ses respectivamente. No se detect&#xf3; evidencia alguna de costos netos de las AMP para las pesquer&#xed;as en ning&#xfa;n lugar. Los beneficios pesqueros incluyen un incremento del stock de peces, el volumen de capturas, las capturas por unidad de esfuerzo, la fecundidad y la exportaci&#xf3;n de larvas; y pescados y langostas m&#xe1;s grandes. Las AMP bien dise&#xf1;adas y aplicadas proporcionan beneficios sostenibles para las comunidades pesqueras. Incluso las AMP mal dise&#xf1;adas pueden proporcionar ventajas econ&#xf3;micas. Las AMP representan una de las mejores estrategias para mantener la explotaci&#xf3;n sostenible de los recursos marinos.</p>
			</trans-abstract>
			<kwd-group>
				<kwd>marine</kwd>
				<kwd>conservation</kwd>
				<kwd>fisheries</kwd>
				<kwd>reserve</kwd>
				<kwd>biodiversity</kwd>
				<kwd>tourism</kwd>
				<kwd>sustainability</kwd>
				<kwd>economic</kwd>
			</kwd-group>
			<kwd-group xml:lang="es">
				<kwd>marino</kwd>
				<kwd>conservaci&#xf3;n</kwd>
				<kwd>pesca</kwd>
				<kwd>reserva</kwd>
				<kwd>biodiversidad</kwd>
				<kwd>turismo</kwd>
				<kwd>sostenibilidad</kwd>
				<kwd>econ&#xf3;mico</kwd>
			</kwd-group>
			<funding-group id="fw-01">
				<award-group id="aw1">
					<funding-source>Horizon Europe</funding-source>
					<award-id>101059988</award-id>
				</award-group>
				<funding-statement>This paper is a contribution to the project MPA Europe funded by Horizon Europe under Grant Agreement 101059988.</funding-statement>
			</funding-group>
			<counts>
				<fig-count count="4"/>
				<table-count count="2"/>
				<equation-count count="0"/>
				<ref-count count="168"/>
				<page-count count="17"/>
			</counts>
		</article-meta>
	</front>
	<body>
		<sec id="sec1" sec-type="intro">
			<title>Introduction</title>
			<p>More than three billion people depend on seafood for almost 20% of their protein and essential nutrients, but two-thirds of fish stocks are in poor biological condition, one-third are overfished, and catch from wild capture fisheries has been steadily declining since 1996 (<xref ref-type="bibr" rid="B29">Costello et al. 2016a</xref>, <xref ref-type="bibr" rid="B128">Pauly and Zeller 2016</xref>, <xref ref-type="bibr" rid="B50">FAO 2019</xref>). With a growing population and increasing anthropogenic pressure on the oceans, future food security for much of the human population is at risk (<xref ref-type="bibr" rid="B166">Worm 2016</xref>, <xref ref-type="bibr" rid="B138">Roberts et al. 2017</xref>). To avert negative effects on food security, changes in fisheries management practices are needed. Optimizing commercial fisheries for long-term profits, rather than annual maximum sustainable yield, could triple economic benefits from fishing and stabilize 97% of fish stocks by 2050 (<xref ref-type="bibr" rid="B29">Costello et al. 2016a</xref>, <xref ref-type="bibr" rid="B166">Worm 2016</xref>).</p>
			<p>In addition to food insecurity, we are in the midst of a biodiversity crisis (<xref ref-type="bibr" rid="B115">McCauley et al. 2015</xref>). The overexploitation of marine wildlife has led to trophic cascades that alter entire ecosystems, and species that were once common are now threatened with extinction (<xref ref-type="bibr" rid="B115">McCauley et al. 2015</xref>). This overexploitation of biodiversity destabilizes ecological and economic ecosystem services, hindering sustainable food provisioning, carbon sequestration and fisheries economic benefits (<xref ref-type="bibr" rid="B33">Costello and Baker 2011</xref>, <xref ref-type="bibr" rid="B115">McCauley et al. 2015</xref>, <xref ref-type="bibr" rid="B49">Epstein et al. 2022</xref>). In light of these challenges, marine protected areas (MPAs) have been used to protect and restore ecosystems and their associated habitats, populations and services (<xref ref-type="bibr" rid="B146">Selig and Bruno 2010</xref>, <xref ref-type="bibr" rid="B98">Leleu et al. 2012</xref>, <xref ref-type="bibr" rid="B94">Krueck et al. 2017</xref>). However, there remains scepticism towards using MPAs to achieve economic benefits, specifically regarding fisheries (<xref ref-type="bibr" rid="B53">Fletcher et al. 2015</xref>, <xref ref-type="bibr" rid="B77">Hilborn 2017</xref>, <xref ref-type="bibr" rid="B78">2018</xref>), as discussed below.</p>
			<p>MPAs are widely accepted as the principal tool for protecting biodiversity and conserving threatened species, but their role in fisheries management is debated (<xref ref-type="bibr" rid="B138">Roberts et al. 2017</xref>, <xref ref-type="bibr" rid="B131">Pendleton et al. 2018</xref>, <xref ref-type="bibr" rid="B93">Kriegl et al. 2021</xref>). Critics argue that evidence of fisheries economic benefits resulting from MPAs are too context-dependent to draw robust conclusions about their efficacy as fisheries management tools (<xref ref-type="bibr" rid="B80">Hilborn et al. 2004</xref>, <xref ref-type="bibr" rid="B43">Di Franco et al. 2016</xref>, <xref ref-type="bibr" rid="B82">Hughes et al. 2016</xref>). They argue that there is a lack of evidence that MPAs benefit fisheries, for example, &#x201c;&#x2026;the evidence that MRs [Marine Reserves] substantially enhance fishery stocks is weak&#x201d; (<xref ref-type="bibr" rid="B24">Caveen et al. 2015</xref>). Critics also argue that evidence of spillover is &#x201c;&#x2026;primarily anecdotal&#x2026;&#x201d; (<xref ref-type="bibr" rid="B127">Pantzar et al. 2018</xref>), that there is &#x201c;no equivocal demonstration of spillover in fisheries adjacent to MPA&#x201d; (<xref ref-type="bibr" rid="B69">Hargreaves-Allen 2020</xref>), and that there is only mixed evidence that MPAs enhance larval export or spillover (<xref ref-type="bibr" rid="B24">Caveen et al. 2015</xref>). Others argue that fishing effort may increase outside the MPAs due to fishery displacement (<xref ref-type="bibr" rid="B79">Hilborn and Hilborn 2019</xref>, <xref ref-type="bibr" rid="B117">McConnaughey et al. 2020</xref>), that MPAs provide few if any socio-economic benefits (<xref ref-type="bibr" rid="B24">Caveen et al. 2015</xref>) and that gear and catch restrictions provide better biodiversity protection than MPAs (<xref ref-type="bibr" rid="B79">Hilborn and Hilborn 2019</xref>, <xref ref-type="bibr" rid="B117">McConnaughey et al. 2020</xref>). Such criticisms are frequently used to devalue the use of MPAs as fisheries management tools, without consideration of the factors that result in such outcomes or the success of MPAs in other settings. In all these cases, the examples to the contrary are regarded as exceptional or it is argued that the evidence supporting MPA benefits is not enough.</p>
			<p>This opposition causes confusion and conflict between stakeholders, slowing progress towards ocean protection targets (<xref ref-type="bibr" rid="B10">Ballantine 2014</xref>, <xref ref-type="bibr" rid="B108">Manson et al. 2021</xref>). As economic benefits, food security and other ecosystem services are underpinned by the restoration and bolstering of ecological functions, failure to adequately protect marine biodiversity and important habitats continues to undermine the economic and environmental sustainability of the world&#x2019;s oceans. Here, the evidence for benefits of MPAs to the economic activities of fisheries and tourism are reviewed. Google Scholar was searched using the key words &#x201c;economic&#x201d;, &#x201c;value&#x201d;, &#x201c;MPA&#x201d;, &#x201c;marine protected area&#x201d;, &#x201c;marine reserve&#x201d;, &#x201c;fisheries&#x201d; and &#x201c;tourism&#x201d;, and the first 200 records were reviewed to find empirical measurements of economic value to fisheries or tourism.</p>
		</sec>
		<sec id="sec2">
			<title>Fisheries</title>
			<p>Of the 51 MPAs reviewed here: 35% (18) were less than 10 years old and 22% (11) were more than 20 years old. Benefits to adjacent finfish, crustacean and mollusc fisheries were detected in 46 (90%) of the MPAs, including an increased fishery catch (76%) and body size (25%), and detection of spillover (16%) (<xref ref-type="table" rid="t1">Table 1</xref>). The latter three percentages exceed 100% because some examples showed several effects. Economic benefits to fisheries from MPAs were reported for 25 countries (plus two overseas territories), spanning temperate, sub-tropical and tropical seas in the North Atlantic, North Pacific, South Pacific and Indian Oceans (<xref ref-type="fig" rid="f1">Fig. 1</xref>). The MPAs encompassed a variety of ecosystems, including coral reefs, kelp forests, mangroves, rocky reefs, salt marshes, mudflats and sandy and muddy seabeds. The MPAs employed a variety of protection methods, including multi-use zones, temporary and permanent closures, restrictions on destructive gear types, and bans on boat-based fishing. The use of no-take MPAs, hereafter called marine reserves, consistently showed the largest benefits.</p>
			<fig id="f1">
				<label>Fig. 1</label>
				<caption>
					<title>Infographic with summarized information of benefits of MPAs for fisheries, as derived from the papers analyzed in this study (<xref ref-type="table" rid="t1">Table 1</xref>).</title>
				</caption>
				<graphic id="gra-1" xlink:href="SCIMAR-88-01-e080-gf1.png"/>
			</fig>
			<table-wrap id="t1">
				<label>Table 1</label>
				<caption>
					<title>Examples of observed benefits to fisheries due to marine protected areas in order of year of publication. Only real-world, non-theoretical examples were included. + increase, * undocumented, ~ no difference in target species abundance (e.g. fishery catch) before-after or inside-outside MPAs, so protection is without cost, **spillover reported (but may be inferred in other cases). MPA age is years established at the time of the study cited.</title>
				</caption>
				<table>
					<colgroup>
						<col/>
						<col/>
						<col/>
						<col/>
						<col/>
						<col/>
					</colgroup>
					<thead>
						<tr>
							<th align="left">Benefits to fisheries</th>
							<th align="left">Location</th>
							<th align="left">Authors</th>
							<th align="center">Age </th>
							<th align="center">Catch</th>
							<th align="center">Body size</th>
						</tr>
					</thead>
					<tbody>
						<tr>
							<td align="left"><bold>Increased fishery catch and body size</bold></td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
						</tr>
						<tr>
							<td align="left">Fishermen noted increased spillover** up to 2 km from MPA, larger catches and larger fish. Increased CPUE and catch per unit area (CPUA).</td>
							<td align="left">Kenya</td>
							<td align="left">
								<xref ref-type="bibr" rid="B116">McClanahan and Mangi 2000</xref>
							</td>
							<td align="center">9</td>
							<td align="center">+ </td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">After 3 years, 5-fold increase in kaikoso clams (<italic>Andarra</italic> spp.) in adjacent fished areas and a 200% increase in CPUE. After 5 years, 7-fold increase.</td>
							<td align="left">Fiji</td>
							<td align="left">
								<xref ref-type="bibr" rid="B152">Tawake et al. 2001</xref>
							</td>
							<td align="center">4</td>
							<td align="center">+</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">CPUE of all fish and CPUE and length of common pandora, <italic>Pagellus erythrinus</italic>, and red mullet, <italic>Mullus surmuletus</italic>, increased close to the reserve boundary.</td>
							<td align="left">Spain</td>
							<td align="left">
								<xref ref-type="bibr" rid="B149">Stelzenm&#xfc;ller et al. 2007</xref>
							</td>
							<td align="center">24</td>
							<td align="center">+</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">Despite high fishing effort, fish yields within 500 m of the MPA increased continuously during the study period. Increased fish size in areas between the reserve and fished zones.</td>
							<td align="left">Spain</td>
							<td align="left">
								<xref ref-type="bibr" rid="B151">Stobart et al. 2009</xref>
							</td>
							<td align="center">19</td>
							<td align="center">+</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">Mean annual net benefit of 10% of the catch in weight for lobster <italic>Palinurus elephas,</italic> despite reserve protection.</td>
							<td align="left">Spain</td>
							<td align="left">
								<xref ref-type="bibr" rid="B64">Go&#xf1;i et al. 2010</xref>
							</td>
							<td align="center">20</td>
							<td align="center">+</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">Increased CPUE inside the periodical closures. Fish larger in catches from closures and Acanthuridae were significantly more abundant. Fish flight initiation distance decreased.</td>
							<td align="left">Vanuatu</td>
							<td align="left">
								<xref ref-type="bibr" rid="B86">Januchowski-Hartley et al. 2014</xref>
							</td>
							<td align="center"> 6</td>
							<td align="center">+</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">Recovery of cod stock following MPA and reduction of fishing effort in wider area.</td>
							<td align="left">Kattegat, Sweden</td>
							<td align="left">
								<xref ref-type="bibr" rid="B16">Bergstr&#xf6;m et al. 2022</xref>
							</td>
							<td align="center">12</td>
							<td align="center">+</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">Increased abundance and size of groupers outside MPA.</td>
							<td align="left">Mediterranean, Israel</td>
							<td align="left">
								<xref ref-type="bibr" rid="B56">Frid et al. 2022</xref>
							</td>
							<td align="center">4</td>
							<td align="center">+</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">
								<bold>Increased fishery catch only</bold>
							</td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
						</tr>
						<tr>
							<td align="left">Both U.S. National Monuments in the Pacific show that catch and CPUE are higher for longline fisheries.</td>
							<td align="left">Hawai&#x2019;i</td>
							<td align="left">
								<xref ref-type="bibr" rid="B106">Lynham et al. 2020</xref>
							</td>
							<td align="center">14</td>
							<td align="center">+</td>
							<td align="center">~</td>
						</tr>
						<tr>
							<td align="left">35% reduction in fishing area compensated by a 225% increase in total catch for spiny lobster (<italic>Panulirus interruptus</italic>) after 6 years.</td>
							<td align="left">NE Pacific, USA</td>
							<td align="left">
								<xref ref-type="bibr" rid="B99">Lenihan et al. 2021</xref>
							</td>
							<td align="center">9</td>
							<td align="center">+</td>
							<td align="center">~</td>
						</tr>
						<tr>
							<td align="left">Spillover** was detected up to 1 km beyond the reserve for small herbivorous fishes (Acanthuridae and Scaridae). Despite concentrated fishing pressure, fish abundance outside the reserve showed no decrease.</td>
							<td align="left">Mozambique</td>
							<td align="left">
								<xref ref-type="bibr" rid="B38">da Silva et al. 2015</xref>
							</td>
							<td align="center">9</td>
							<td align="center">+ </td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Fishermen claim higher catch in fishing grounds adjacent to the MPA and fish close to the MPA boundary. Increased CPUE on nearby fishing grounds.</td>
							<td align="left">Isle of Man</td>
							<td align="left">
								<xref ref-type="bibr" rid="B18">Bradshaw et al. 1999</xref>
							</td>
							<td align="center">10</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Increase in target fish in adjacent fishing grounds. Increase in catch rates.</td>
							<td align="left">Madagascar</td>
							<td align="left">
								<xref ref-type="bibr" rid="B66">Grandcourt et al. 2001</xref>
							</td>
							<td align="center">12</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">In adjacent areas after 5 years catches increased by 46% - 90%, depending on fishing gear, and biomass of commercial reef fish doubled.</td>
							<td align="left">Saint Lucia</td>
							<td align="left">
								<xref ref-type="bibr" rid="B136">Roberts et al. 2001</xref>
							</td>
							<td align="center">6</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Two-thirds increase in CPUE in adjacent fishery grounds, fishery now sustainable.</td>
							<td align="left">Red Sea, Egypt,</td>
							<td align="left">
								<xref ref-type="bibr" rid="B60">Galal et al. 2002</xref>
							</td>
							<td align="center">7</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Landing volumes of snow crabs (<italic>Chionoecetes opilio</italic>) increased from 59 t in 1980 to 196 t in 1999. CPUE increased more than 4-fold.</td>
							<td align="left">Japan</td>
							<td align="left">
								<xref ref-type="bibr" rid="B167">Yamasaki 2002</xref>
							</td>
							<td align="center">19</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Increased CPUE for hogfish (<italic>Lachnolaimus maximus)</italic> related to decreasing distance from the reserve.</td>
							<td align="left">Turks and Caicos Islands</td>
							<td align="left">
								<xref ref-type="bibr" rid="B157">Tupper and Rudd 2002</xref>
							</td>
							<td align="center">10</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Ten-fold increase in fish catch by weight and ten-fold increase in CPUE for line fishing since reserve creation.</td>
							<td align="left">Philippines</td>
							<td align="left">
								<xref ref-type="bibr" rid="B114">Maypa et al. 2002</xref>
							</td>
							<td align="center"> 20</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Catch rates of trammel netters were 33% - 50% higher inside the trawl exclusion area than outside.</td>
							<td align="left">Italy</td>
							<td align="left">
								<xref ref-type="bibr" rid="B164">Whitmarsh et al. 2002</xref>
							</td>
							<td align="center">12</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Increased catch after 5 years for commercial species. Increased CPUE and double total catch for cod (<italic>Gadus morhua</italic>). Increased larval export from scallops (<italic>Placopecten magellanicus</italic>).</td>
							<td align="left">Atlantic USA</td>
							<td align="left">
								<xref ref-type="bibr" rid="B61">Gell and Roberts 2003</xref>
							</td>
							<td align="center">9</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Biomass of bignose unicorn fish (<italic>Naso vlamingii</italic>) increased by a factor of 40 outside the reserve (200 - 250 m). Hook-and-line CPUE for <italic>N. vlamingii</italic> 45 times higher within 200 m of the reserve.</td>
							<td align="left">Philippines</td>
							<td align="left">
								<xref ref-type="bibr" rid="B140">Russ et al. 2003</xref>
							</td>
							<td align="center"> 20</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Increasing CPUE near the MPA for 4 km, declining as increasing distance from the MPA, including spillover**.</td>
							<td align="left">Atlantic USA</td>
							<td align="left">
								<xref ref-type="bibr" rid="B121">Murawski et al. 2004</xref>, <xref ref-type="bibr" rid="B122">2005</xref></td>
							<td align="center"> 10</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Catches increased by 27% outside the Sumolin reserve and 41% outside the Apo reserve. Total fishery catch either sustained or enhanced.</td>
							<td align="left">Philippines</td>
							<td align="left">
								<xref ref-type="bibr" rid="B4">Alcala et al. 2005</xref>
							</td>
							<td align="center">31</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Increasing lobster CPUE and CPUA within 2 km of MPA.</td>
							<td align="left">Spain</td>
							<td align="left">
								<xref ref-type="bibr" rid="B62">Go&#xf1;i et al. 2006</xref>
							</td>
							<td align="center">16</td>
							<td align="center">+ </td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Catch rates higher near the reserve by a factor of 1.1 - 2.0.</td>
							<td align="left">Philippines</td>
							<td align="left">
								<xref ref-type="bibr" rid="B2">Abesamis et al. 2006</xref>
							</td>
							<td align="center">23</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Increased spillover** beyond MPA boundaries for 2.5 km.</td>
							<td align="left">France, Spain</td>
							<td align="left">
								<xref ref-type="bibr" rid="B63">Go&#xf1;i et al. 2008</xref>
							</td>
							<td align="center">8</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">A general pattern of decreasing fish biomass from within MPA to fished areas consistent with biomass spillover.</td>
							<td align="left">France, Spain</td>
							<td align="left">
								<xref ref-type="bibr" rid="B70">Harmelin-Vivien et al. 2008</xref>
							</td>
							<td align="center">34</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Increased CPUE and IPUE (income per unit effort) close to the MPA border. Increased resilience of fish assemblages against fishing and human impacts within 2 km.</td>
							<td align="left">Spain</td>
							<td align="left">
								<xref ref-type="bibr" rid="B150">Stelzenm&#xfc;ller et al. 2008</xref>
							</td>
							<td align="center"> 34</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Three-fold increase in the density of mollusc juveniles, black murex snail (<italic>Hexaplex nigritus</italic>), found in fished areas at the downstream edge of the reserve.</td>
							<td align="left">Mexico</td>
							<td align="left">
								<xref ref-type="bibr" rid="B36">Cudney-Bueno et al. 2009</xref>
							</td>
							<td align="center">7</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Five-fold increase in yellow tang (<italic>Zebrasoma flavescens</italic>) within MPAs. Density in boundary sites less than 1 km from the nearest MPA nearly as high as within the MPA.</td>
							<td align="left">Hawai&#x2019;i</td>
							<td align="left">
								<xref ref-type="bibr" rid="B165">Williams et al. 2009</xref>
							</td>
							<td align="center">10</td>
							<td align="center">+ </td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Higher fishery yields within 500 m of the MPA than in areas more than 1 km away.</td>
							<td align="left">France, Spain</td>
							<td align="left">
								<xref ref-type="bibr" rid="B54">Forcada et al. 2009</xref>
							</td>
							<td align="center">20</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">CPUE of target species and marketable catch increased by 2% - 4% per year, over at least 30 years.</td>
							<td align="left">Southern Europe</td>
							<td align="left">
								<xref ref-type="bibr" rid="B159">Vandeperre et al. 2011</xref>
							</td>
							<td align="center"> 37</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Reserves covering 28% of the local reef area produced half of all juvenile recruitment to fished reefs within 30 km.</td>
							<td align="left">Australia</td>
							<td align="left">
								<xref ref-type="bibr" rid="B71">Harrison et al. 2012</xref>
							</td>
							<td align="center"> 19</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Reduced fish flight initiation distance, increased CPUE.</td>
							<td align="left">Philippines</td>
							<td align="left">
								<xref ref-type="bibr" rid="B85">Januchowski-Hartley et al. 2013</xref>
							</td>
							<td align="center"> 29</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">CPUE in the MPA vicinity immediately increased. This continued after 5 years, doubling pre-MPA CPUE after 10 years.</td>
							<td align="left">South Africa</td>
							<td align="left">
								<xref ref-type="bibr" rid="B90">Kerwath et al. 2013</xref>
							</td>
							<td align="center">23</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Density of adult king scallops (<italic>Pecten maximus</italic>) declined three-fold with increasing distance from the reserve boundary.</td>
							<td align="left">U.K.</td>
							<td align="left">
								<xref ref-type="bibr" rid="B81">Howarth et al. 2015</xref>
							</td>
							<td align="center">7</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Adult snapper (<italic>Pagrus auratus</italic>) within the MPA contributed 11% of juveniles to surrounding areas with no decreasing trend up to 40 km away.</td>
							<td align="left">New Zealand</td>
							<td align="left">
								<xref ref-type="bibr" rid="B97">Le Port et al. 2017</xref>
							</td>
							<td align="center">37</td>
							<td align="center">+ </td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Relative abundance of snapper (<italic>Pagrus auratus</italic>) increased within the MPA despite increased fishing effort.</td>
							<td align="left">Australia</td>
							<td align="left">
								<xref ref-type="bibr" rid="B68">Harasti et al. 2018</xref>
							</td>
							<td align="center">13</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Increased diversity of rockfish larvae in plankton.</td>
							<td align="left">California, USA</td>
							<td align="left">
								<xref ref-type="bibr" rid="B55">Freeman et al. 2022</xref>
							</td>
							<td align="center">12</td>
							<td align="center">+</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">
								<bold>Increased body size only</bold>
							</td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
						</tr>
						<tr>
							<td align="left">Larger spiny lobsters (<italic>Jasus edwardsii</italic>) were caught adjacent to the reserve.</td>
							<td align="left">New Zealand</td>
							<td align="left">
								<xref ref-type="bibr" rid="B89">Kelly et al. 2002</xref>
							</td>
							<td align="center">27</td>
							<td align="center">~</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">Lobsters spillover** from MPAs were larger </td>
							<td align="left">North Sea, Norway</td>
							<td align="left">
								<xref ref-type="bibr" rid="B155">Thorbj&#xf8;rnsen et al. 2018</xref>
							</td>
							<td align="center">9</td>
							<td align="center">~</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">Average size of red hind grouper (<italic>Epinephelus guttatus</italic>) increased by 34%. Sex ratio decreased to 4 females per male.</td>
							<td align="left">Virgin Islands USA</td>
							<td align="left">
								<xref ref-type="bibr" rid="B15">Beets and Friedlander 1999</xref>
							</td>
							<td align="center">9</td>
							<td align="center">*</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">Record-size catches of red drum (<italic>Sciaenops ocellatus</italic>), black drum (<italic>Pogonias cromis</italic>) and spotted sea trout (C<italic>ynoscion nebulosus</italic>) in adjacent areas to the reserve.</td>
							<td align="left">Atlantic USA</td>
							<td align="left">
								<xref ref-type="bibr" rid="B136">Roberts et al. 2001</xref>
							</td>
							<td align="center">41</td>
							<td align="center">*</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">Spillover**, density and modal size of <italic>N. vlamingii</italic> increased outside the reserve within 200 - 300 m.</td>
							<td align="left">Philippines</td>
							<td align="left">
								<xref ref-type="bibr" rid="B1">Abesamis and Russ 2005</xref>
							</td>
							<td align="center">22</td>
							<td align="center">*</td>
							<td align="center">+</td>
						</tr>
						<tr>
							<td align="left">
								<bold>Spillover reported only</bold>
							</td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
						</tr>
						<tr>
							<td align="left">Spillover** of finfish species between the closed area and fished area with time lags ranging from 1 - 3 years.</td>
							<td align="left">Atlantic Canada</td>
							<td align="left">
								<xref ref-type="bibr" rid="B52">Fisher and Frank 2002</xref>
							</td>
							<td align="center"> 15</td>
							<td align="center">*</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Larval export** from the mussel, <italic>Perna perna,</italic> increased from reserves, enhancing recruitment in nearby fished areas within several km.</td>
							<td align="left">South Africa</td>
							<td align="left">
								<xref ref-type="bibr" rid="B129">Pelc et al. 2009</xref>
							</td>
							<td align="center"> 34</td>
							<td align="center">*</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left"><bold>Uncertain effect on fisheries</bold></td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
							<td align="left"> </td>
						</tr>
						<tr>
							<td align="left">Fishing activity decreased by 82% in the MPA without any negative effect on the industrial pelagic fishery catch in the region. </td>
							<td align="left">Mexican Pacific</td>
							<td align="left">
								<xref ref-type="bibr" rid="B51">Favoretto et al. 2023</xref>
							</td>
							<td align="center">5</td>
							<td align="center">~</td>
							<td align="center">~</td>
						</tr>
						<tr>
							<td align="left">36% decline in catch after closure of 33% of the area to fishing but no decline in CPUE.</td>
							<td align="left">Great Barrier Reef, Australia</td>
							<td align="left">
								<xref ref-type="bibr" rid="B53">Fletcher et al. 2015</xref>
							</td>
							<td align="center">9</td>
							<td align="center">~</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">The majority of fishermen (85%) perceived no effect of marine reserves on their catch.</td>
							<td align="left">Seychelles</td>
							<td align="left">
								<xref ref-type="bibr" rid="B27">Cinner et al. 2014</xref>
							</td>
							<td align="center"> 46</td>
							<td align="center">*</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Since MPA designation, 23% of recreational fishermen felt the number of fish caught had improved, 32% considered it the same, 17% felt it had declined and 28% could not say.</td>
							<td align="left">Australia</td>
							<td align="left">
								<xref ref-type="bibr" rid="B112">Martin et al. 2016</xref>
							</td>
							<td align="center">9</td>
							<td align="center">*</td>
							<td align="center">*</td>
						</tr>
						<tr>
							<td align="left">Initial analysis of a decline of fish catch of 14% not supported by second analysis.</td>
							<td align="left">Gulf of Mexico, USA</td>
							<td align="left">
								<xref ref-type="bibr" rid="B147">Smith et al. 2006</xref>, <xref ref-type="bibr" rid="B148">2007</xref></td>
							<td align="center">4.5</td>
							<td align="center">-</td>
							<td align="center">*</td>
						</tr>
					</tbody>
				</table>
			</table-wrap>
			<p>Fisheries close to MPAs had up to 45 times higher catch per unit effort (CPUE) and 40 times higher catch (<xref ref-type="table" rid="t1">Table 1</xref>). In one study, fish size was on average 34% greater (<xref ref-type="bibr" rid="B15">Beets and Friedlander 1999</xref>), and in another, larval export was enhanced, showing no decreasing trend up to 40 km away (<xref ref-type="bibr" rid="B97">Le Port et al. 2017</xref>) (<xref ref-type="table" rid="t1">Table 1</xref>). Another MPA showed increased juvenile recruitment up to the 1000 km<sup>2</sup> limit of the study area (<xref ref-type="bibr" rid="B71">Harrison et al. 2012</xref>). Fish behaviour can also change, with reduced flight initiation distance within an MPA, making fish easier to observe (<xref ref-type="bibr" rid="B30">Costello 2014</xref>, <xref ref-type="bibr" rid="B86">Januchowski-Hartley et al. 2014</xref>). MPAs with varying levels of protection, such as those with partial protection areas and fishery closures, also provided significant advantages for fishermen. <xref ref-type="bibr" rid="B167">Yamasaki et al. (2002)</xref> found a nearly four-fold increase in catch volumes, <xref ref-type="bibr" rid="B159">Vandeperre et al. (2011)</xref> showed an increase in CPUE of 60%-120% over 30 years, and <xref ref-type="bibr" rid="B90">Kerwath et al. (2013)</xref> found that CPUE doubled after 10 years of protection. In total, 32 studies found increased catch since MPA designation.</p>
			<p>Some of the studies indicated economic advantages to fisheries due to increases in fish size and consequently fecundity (<xref ref-type="table" rid="t1">Table 1</xref>), as predicted by modelling (<xref ref-type="bibr" rid="B39">De Leo and Micheli 2015</xref>). These outcomes may not directly contribute to catch increases, but they help sustain fisheries adjacent to protected areas within larval dispersal ranges through increased recruitment, as, for example, was found by <xref ref-type="bibr" rid="B97">Le Port et al. (2017)</xref>, <xref ref-type="bibr" rid="B129">Pelc et al. (2009</xref>, <xref ref-type="bibr" rid="B130">2010)</xref> and <xref ref-type="bibr" rid="B71">Harrison et al. (2012)</xref>. <xref ref-type="bibr" rid="B71">Harrison et al. (2012)</xref> showed that reserves covering only 28% of the reef area produced half of all juvenile recruitment, and <xref ref-type="bibr" rid="B97">Le Port et al. (2017)</xref> found that a small (5.2 km<sup>2</sup>) reserve in New Zealand contributed over 10% of juveniles to the surrounding areas. <xref ref-type="bibr" rid="B134">Qu et al. (2021)</xref> valued the snapper recruitment effect noted by <xref ref-type="bibr" rid="B97">Le Port et al. (2017)</xref> at NZ$5 million (~US$ 3 million) per annum to local commercial fisheries, a considerable economic gain.</p>
			<p>Two studies did not find unambiguous fishery benefits in terms of gross fish catch. In one case the MPA was only established for 4.5 years and a decline in catch could have been due to one year of poor recruitment of one fish species (<xref ref-type="bibr" rid="B147">Smith et al. 2006</xref>), and the same authors subsequently found no decline in catch (<xref ref-type="bibr" rid="B148">Smith et al. 2007</xref>). In the second, there was stable or increasing CPUE following closure of 33% of the area to fishing and a 30% decline in effort due to 6 of 13 fisheries having some licences bought out (<xref ref-type="bibr" rid="B53">Fletcher et al. 2015</xref>). However, this study did not account for management measures to reduce overfishing and geographic variation in fisheries and environmental factors (e.g. typhoons) (<xref ref-type="bibr" rid="B82">Hughes et al. 2016</xref>). Because it was surprising that no studies showed losses to fisheries, we also polled the ~12000 members of the &#x201c;MPA Help&#x201d; community (<ext-link ext-link-type="uri" xlink:href="https://octogroup.org/programs/mpa-help/">https://octogroup.org/programs/mpa-help/</ext-link>) for evidence of such costs. However, the responses could only provide studies on estimated or modelled costs to fisheries, not demonstrated costs. This indicates that if there are any net losses to any fisheries anywhere due to MPAs, they are not documented and likely rare.</p>
			<p>There are many reasons why a fishery may or may not benefit from an MPA, as reviewed by <xref ref-type="bibr" rid="B59">Gaines et al. (2010)</xref>. Inverted logic often regards MPAs as the &#x201c;experiment&#x201d; or &#x201c;treatment&#x201d;, whereas the MPA is the control or reference for the experiments that study the human activities outside it. Comparisons inside and outside indicate two-thirds of rocky and coral reef fishes have been removed by fishing (<xref ref-type="bibr" rid="B47">Edgar et al. 2014</xref>). Assessing the impact of these activities requires not only controls (i.e. marine reserves), but data on their direct effect on target species abundance, body size and age, and associated habitat damage, plus data on the indirect effects on food webs, such as those due to trophic cascades (<xref ref-type="bibr" rid="B35">Costello et al. 2022</xref>). The variation associated with fishing effort and gear makes quantifying fishing impact and CPUE difficult (<xref ref-type="bibr" rid="B147">Smith et al. 2006</xref>). Furthermore, fishing pressure-be it commercial, recreational, or both-may often continue to increase in the area regardless of the establishment of an MPA (e.g. <xref ref-type="bibr" rid="B123">Nillos Kleiven et al. 2019</xref>, <xref ref-type="bibr" rid="B91">Kleiven 2020</xref>, <xref ref-type="bibr" rid="B96">LaScala-Gruenewald et al. 2021</xref>).</p>
		</sec>
		<sec id="sec3">
			<title>Tourism</title>
			<p>Examples of economic benefits from tourism were found in 24 countries, of which the majority were in tropical and sub-tropical locations, with four examples from temperate regions (France, Spain, Italy and New Zealand) (<xref ref-type="fig" rid="f2">Fig. 2</xref>, <xref ref-type="table" rid="t2">Table 2</xref>). MPAs were located in the Indian, North Atlantic, North Pacific and South Pacific Oceans. The MPAs which showed benefits to tourism covered a range of ecosystems, including rocky reefs, kelp forests and sandy and muddy bottoms, but were dominated by coral reefs, mangroves and seagrass ecosystems. These 31 examples (<xref ref-type="table" rid="t2">Table 2</xref>) reported benefits to local communities, private companies, NGOs and state departments directly through user fees and indirectly through related expenses such as transport, accommodation, food, goods and other services. Shark diving contributes about $18 million a year to the economy of Palau, whereas shark fishing would contribute only $10800 (<xref ref-type="bibr" rid="B161">Vianna et al. 2012</xref>).</p>
			<fig id="f2">
				<label>Fig. 2</label>
				<caption>
					<title>Infographic with summarized highlights of economic benefits of MPAs for tourism, as derived from the papers analyzed in this study (<xref ref-type="table" rid="t2">Table 2</xref>).</title>
				</caption>
				<graphic id="gra-2" xlink:href="SCIMAR-88-01-e080-gf2.png"/>
			</fig>
			<table-wrap id="t2">
				<label>Table 2</label>
				<caption>
					<title>Examples of the economic benefits of marine protected areas from tourism. Only real-world, non-theoretical examples were included. MPA age is years at the time of the study cited.</title>
				</caption>
				<table>
					<colgroup>
						<col/>
						<col/>
						<col/>
						<col/>
					</colgroup>
					<thead>
						<tr>
							<th align="left">Location</th>
							<th align="left">Author(s)</th>
							<th align="left">Benefits from tourism</th>
							<th align="center">Age </th>
						</tr>
					</thead>
					<tbody>
						<tr>
							<td align="left">Cuba</td>
							<td align="left">
								<xref ref-type="bibr" rid="B65">Gonz&#xe1;lez-Sans&#xf3;n et al. 2002</xref>
							</td>
							<td align="left">Punta Frances MPA generates US$200000 per year from SCUBA diving and cruise ship tourism.</td>
							<td align="center">23</td>
						</tr>
						<tr>
							<td align="left">Belize</td>
							<td align="left">
								<xref ref-type="bibr" rid="B168">Young and Bilgre 2002</xref>
							</td>
							<td align="left">From 1995 to 1999, the Hol Chan Marine Reserve generated almost US$500000 from ticket sales.</td>
							<td align="center">15</td>
						</tr>
						<tr>
							<td align="left">Seychelles</td>
							<td align="left">
								<xref ref-type="bibr" rid="B113">Mathieu et al. 2003</xref>
							</td>
							<td align="left">Seychelles MPAs provide direct revenue of US$135324, providing 31 jobs, 13 trainee positions and an operating profit of over US$8000.</td>
							<td align="center">&#x2265; 6</td>
						</tr>
						<tr>
							<td align="left">Philippines</td>
							<td align="left">
								<xref ref-type="bibr" rid="B156">Tongson and Dygico 2004</xref>
							</td>
							<td align="left">Tubbataha Reefs Natural Marine Park introduced a fee collection and permit system raising over US$30000 per annum.</td>
							<td align="center">16</td>
						</tr>
						<tr>
							<td align="left">Canary Islands, Spain </td>
							<td align="left">
								<xref ref-type="bibr" rid="B139">Roncin et al. 2008</xref>
							</td>
							<td align="left">La Restinga MPA generates US$739200 from diving-related tourism alone.</td>
							<td align="center">12</td>
						</tr>
						<tr>
							<td align="left">France, Mediterranean</td>
							<td align="left">
								<xref ref-type="bibr" rid="B139">Roncin et al. 2008</xref>
							</td>
							<td align="left">Bonifacio generates US$1137600 from diving-related tourism alone.</td>
							<td align="center">9</td>
						</tr>
						<tr>
							<td align="left">Thailand</td>
							<td align="left">
								<xref ref-type="bibr" rid="B7">Asafu-Adjaye and Tapsuwan 2008</xref>
							</td>
							<td align="left">Mu Ko Similan Marine National Park generates US$457389 annually through entrance fees, service charges and accommodation.</td>
							<td align="center">26</td>
						</tr>
						<tr>
							<td align="left">Sabah, Malaysia</td>
							<td align="left">
								<xref ref-type="bibr" rid="B153">Teh et al. 2008</xref>
							</td>
							<td align="left">Sugud Islands Marine Conservation Area generated US$26900 in 2004 from conservation and user fees.</td>
							<td align="center">7</td>
						</tr>
						<tr>
							<td align="left">South Africa</td>
							<td align="left">
								<xref ref-type="bibr" rid="B46">Dicken and Hosking 2009</xref>
							</td>
							<td align="left">Annual value of tiger shark diving to the Aliwal Shoal region was US$885000 per annum.</td>
							<td align="center">18</td>
						</tr>
						<tr>
							<td align="left">Kenya</td>
							<td align="left">
								<xref ref-type="bibr" rid="B74">Hicks et al. 2009</xref>
							</td>
							<td align="left">Mombasa Marine National Park valued solely for recreation (tourism) at US$3.5 million/km/year.</td>
							<td align="center">18</td>
						</tr>
						<tr>
							<td align="left">Spain</td>
							<td align="left">
								<xref ref-type="bibr" rid="B118">Merino et al. 2009</xref>
							</td>
							<td align="left">Medes Islands Marine Reserve generates US$7.4 million annually through non-extractive tourism.</td>
							<td align="center">19</td>
						</tr>
						<tr>
							<td align="left">Australia, Indian Ocean</td>
							<td align="left">
								<xref ref-type="bibr" rid="B23">Catlin et al. 2010</xref>
							</td>
							<td align="left">Annual expenditure of US$4.5 million from whale shark tour participants in the Ningaloo Marine Park.</td>
							<td align="center">23</td>
						</tr>
						<tr>
							<td align="left">Fiji</td>
							<td align="left">
								<xref ref-type="bibr" rid="B20">Brunnschweiler 2010</xref>
							</td>
							<td align="left">Marine park levy from tourism paid annually to each village for not fishing the reserve: US$20000.</td>
							<td align="center">7</td>
						</tr>
						<tr>
							<td align="left">South Africa</td>
							<td align="left">
								<xref ref-type="bibr" rid="B45">Dicken 2010</xref>
							</td>
							<td align="left">Pondoland MPA has a direct value of tourism estimated at US$765800.</td>
							<td align="center">6</td>
						</tr>
						<tr>
							<td align="left">Bonaire, Caribbean</td>
							<td align="left">
								<xref ref-type="bibr" rid="B158">Uyarra et al. 2010</xref>
							</td>
							<td align="left">User fees for access to Bonaire National Marine Park generated US$1039597 in 2008 alone.</td>
							<td align="center">29</td>
						</tr>
						<tr>
							<td align="left">Mexico, Gulf of California</td>
							<td align="left">
								<xref ref-type="bibr" rid="B3">Aburto-Oropeza et al. 2011</xref>
							</td>
							<td align="left">Small-scale tourism operators (less than 30 people) in Cabo Pulmo National Park generated US$538800 in 2006.</td>
							<td align="center">16</td>
						</tr>
						<tr>
							<td align="left">Egypt, Red Sea</td>
							<td align="left">
								<xref ref-type="bibr" rid="B144">Samy and Lizaso 2011</xref>
							</td>
							<td align="left">Ras Mohammed National Park generates US$2635200 per year from user fees. Recreational value of the MPA&#x2019;s coral reefs is between US$153 and US$205 million annually.</td>
							<td align="center">28</td>
						</tr>
						<tr>
							<td align="left">Egypt, Red Sea</td>
							<td align="left">
								<xref ref-type="bibr" rid="B144">Samy and Lizaso 2011</xref>
							</td>
							<td align="left">Wadi El Gemal-Hamata MPA provides 50 jobs and generates US$3995453 per year from fees, penalties and sanctions.</td>
							<td align="center">8</td>
						</tr>
						<tr>
							<td align="left">Fiji</td>
							<td align="left">
								<xref ref-type="bibr" rid="B160">Vianna, et al. 2011</xref>
							</td>
							<td align="left">MPAs generate US$650000 annually from dive tourism by businesses operating at shark diving sites.</td>
							<td align="center">14</td>
						</tr>
						<tr>
							<td align="left">Cocos Island, Costa Rica</td>
							<td align="left">
								<xref ref-type="bibr" rid="B57">Friedlander et al. 2012</xref>
							</td>
							<td align="left">At full occupancy, 5 diving liveaboards at Isla del Coco National Park bring over US$7 million to the local economy.</td>
							<td align="center">34</td>
						</tr>
						<tr>
							<td align="left">Australia, Coral Sea</td>
							<td align="left">
								<xref ref-type="bibr" rid="B40">Deloitte Access Economics 2013</xref>
							</td>
							<td align="left">Value of tourism at the Great Barrier Reef Marine Park estimated at US$6.4 billion annually.</td>
							<td align="center">38</td>
						</tr>
						<tr>
							<td align="left">Zanzibar, Tanzania</td>
							<td align="left">
								<xref ref-type="bibr" rid="B124">Nordlund et al. 2013</xref>
							</td>
							<td align="left">The Chumbe Island Coral Park provides 43 jobs and generates an annual revenue of around US$500000.</td>
							<td align="center">22</td>
						</tr>
						<tr>
							<td align="left">Kenya</td>
							<td align="left">
								<xref ref-type="bibr" rid="B87">Job and Paesler 2013</xref>
							</td>
							<td align="left">Kisite Marine National Park earns annual revenue of US$80000, 7 times its operating costs, and provides 40 jobs.</td>
							<td align="center">40</td>
						</tr>
						<tr>
							<td align="left">Maldives</td>
							<td align="left">
								<xref ref-type="bibr" rid="B21">Cagua et al. 2014</xref>
							</td>
							<td align="left">Annual expenditure of US$7.6 - 9.4 million from whale shark-related tourism.</td>
							<td align="center">5</td>
						</tr>
						<tr>
							<td align="left">New Zealand</td>
							<td align="left">
								<xref ref-type="bibr" rid="B30">Costello 2014</xref>
							</td>
							<td align="left">Ecotourism from Leigh Marine Reserve (Cape Rodney-Okakari Point Marine Reserve) valued at US$5.9 million annually.</td>
							<td align="center">39</td>
						</tr>
						<tr>
							<td align="left">Gal&#xe1;pagos</td>
							<td align="left">
								<xref ref-type="bibr" rid="B105">Lynham et al. 2015</xref>
							</td>
							<td align="left">Marine-based tourism at the Gal&#xe1;pagos Marine Reserve contributes US$236 million annually.</td>
							<td align="center">17</td>
						</tr>
						<tr>
							<td align="left">Thailand</td>
							<td align="left">
								<xref ref-type="bibr" rid="B145">Seenprachawong 2016</xref>
							</td>
							<td align="left">Mu Ko Phi Phi Marine National Park generates large ecotourism benefits representing an annual value in excess of US$200 million.</td>
							<td align="center">33</td>
						</tr>
						<tr>
							<td align="left">Raja Ampat, Indonesia</td>
							<td align="left">
								<xref ref-type="bibr" rid="B8">Atmodjo et al. 2017</xref>
							</td>
							<td align="left">Tourism pays for the costs of managing the Raja Ampat MPA network and provides US$127500 per year to a community fund.</td>
							<td align="center">13</td>
						</tr>
						<tr>
							<td align="left">Bahamas</td>
							<td align="left">
								<xref ref-type="bibr" rid="B67">Haas et al. 2017</xref>
							</td>
							<td align="left">Shark-diving industry contributes US$113.8 million annually to the Bahamian economy in direct and value added expenditures.</td>
							<td align="center">6</td>
						</tr>
						<tr>
							<td align="left">Italy</td>
							<td align="left">
								<xref ref-type="bibr" rid="B103">Lucrezi et al. 2017</xref>
							</td>
							<td align="left">Dive operators in Portofino MPA contribute over US$100000 in tax annually.</td>
							<td align="center">18</td>
						</tr>
						<tr>
							<td align="left">Moalboal, Philippines</td>
							<td align="left">
								<xref ref-type="bibr" rid="B37">Cusack et al. 2021</xref>
							</td>
							<td align="left">Annual revenues directly related to marine reserve visitation estimated at US$4.68 million.</td>
							<td align="center">34</td>
						</tr>
					</tbody>
				</table>
			</table-wrap>
			<p>Individual MPAs have generated millions to billions of dollars in tourism revenue per year (<xref ref-type="table" rid="t2">Table 2</xref>). The Great Barrier Reef Marine Park, Australia, generated US$6.4 billion, and others generate hundreds of millions, such as the Gal&#xe1;pagos Marine Reserve, Ecuador; Mu Ko Phi Phi Marine National Park, Thailand; and Ras Mohammed National Park, Red Sea, Egypt (<xref ref-type="table" rid="t2">Table 2</xref>). Some MPAs obtained millions of dollars from user fees (including fines), including Bonaire National Marine Park, Ras Mohammed National Park and the Wadi El Gemal-Hamata Protected Area. However, many MPAs, such as in New Zealand, do not charge visitor fees. In general, older, more established MPAs provided higher total tourism revenues, as shown by the Great Barrier Reef Marine Park, Mu Ko Phi Phi Marine National Park, Ras Mohammed National Park and the Bahamas Shark Sanctuary (<xref ref-type="fig" rid="f3">Fig. 3</xref>).</p>
			<fig id="f3">
				<label>Fig. 3</label>
				<caption>
					<title>MPAs with known value of combined indirect and direct tourism (US$ on a log10 scale) and their age (Rank Spearman correlation coefficient 0.47, without highest value is 0.33; Pearson Product moment correlation coefficient R2=78). Where an MPA value was estimated as a range between two numbers (e.g. Ras Mohammed National Park, US$153-205 million per annum), the mean of those numbers was used.</title>
				</caption>
				<graphic id="gra-3" xlink:href="SCIMAR-88-01-e080-gf3.png"/>
			</fig>
		</sec>
		<sec id="sec4">
			<title>Rebuttal of criticisms</title>
			<p>Any sample of the literature will be biased by what research was conducted and published. However, the present sample provides no indications of significant costs to fisheries from establishment of MPAs. Indeed, there were 77 examples of evidence that MPAs can provide economic benefits through fisheries (n=46) and tourism (n=31) (<xref ref-type="table" rid="t1">Tables 1</xref>, <xref ref-type="table" rid="t2">2</xref>). Evidently, MPAs can provide a rare win-win strategy for ocean management, enabling conservation and long-term economic goals to be achieved simultaneously.</p>
			<p>Contrary to <xref ref-type="bibr" rid="B24">Caveen et al. (2015)</xref>, MPAs are shown to substantially increase fish stocks and catch, and can provide sometimes lucrative socio-economic benefits (<xref ref-type="table" rid="t1">Tables 1</xref>, <xref ref-type="table" rid="t2">2</xref>, <xref ref-type="fig" rid="f3">Fig. 3</xref>). Another criticism of MPAs is that there is little or only anecdotal evidence of spillover from MPAs to fished areas (<xref ref-type="bibr" rid="B24">Caveen et al. 2015</xref>, <xref ref-type="bibr" rid="B127">Pantzar et al. 2018</xref>, <xref ref-type="bibr" rid="B69">Hargreaves-Allen 2020</xref>). However, the majority of examples of fisheries benefits from MPAs refer to the spillover of target species.</p>
			<p>It may seem counter-intuitive that restricting fishing in an area will result in more fish elsewhere. Yet, this happens because marine life disperses from its safe haven (the MPA), which acts like a reservoir to replenish adjacent fisheries. In financial terms, the capital is invested and people benefit from the interest on the investment. To count MPAs as a cost to fisheries is analogous to claiming that interest earned on money is a cost. The evidence of this benefit is unequivocal (<xref ref-type="table" rid="t1">Table 1</xref>).</p>
			<p>The fact that there were only four examples of larval export from MPAs reflects the practical challenges in distinguishing eggs and larvae from MPA vs non-MPA parents. Nevertheless, rather than there being mixed evidence of enhanced larval export (as suggested by <xref ref-type="bibr" rid="B24">Caveen et al. 2015</xref>), the evidence collected to date supports predictions that MPAs contribute disproportionately to larval dispersal (<xref ref-type="bibr" rid="B130">Pelc et al. 2010</xref>, <xref ref-type="bibr" rid="B55">Freeman et al. 2022</xref>) and subsequent fisheries recruitment, further enhancing their worth as fisheries management tools (<xref ref-type="bibr" rid="B72">Hastings and Botsford 1999</xref>, <xref ref-type="bibr" rid="B39">De Leo and Micheli 2015</xref>, <xref ref-type="bibr" rid="B92">Kough et al. 2019</xref>).</p>
			<p>Recent research has elaborated further on the fisheries benefits of protected areas arising frorm the increased size and fecundity of fish within MPA boundaries (<xref ref-type="bibr" rid="B14">Barneche et al. 2019</xref>, <xref ref-type="bibr" rid="B111">Marshall et al. 2021</xref>). As protected areas increase fish size by an average of 28% (<xref ref-type="bibr" rid="B101">Lester et al. 2009</xref>), and the reproductive output of fish increases disproportionately with size and weight, the reproductive contribution of fish within protected areas has been systematically underestimated (<xref ref-type="bibr" rid="B110">Marshall et al. 2019</xref>). Consequently, establishing protected reservoirs of Big Old Fat Fecund Female Fish (BOFFFFs) can lead to increased larvae diversity in the plankton (<xref ref-type="bibr" rid="B55">Freeman et al. 2022</xref>) and enhance fishery yields (<xref ref-type="bibr" rid="B110">Marshall et al. 2019</xref>). This highlights the importance of keeping BOFFFFs within breeding populations and also shows that a failure to consider reproductive hyper-allometry overestimates the effectiveness of traditional fisheries management (<xref ref-type="bibr" rid="B111">Marshall et al. 2021</xref>). These important findings further strengthen the use of protected areas as fisheries management tools. Were fishermen given custodianship of fish stocks, then, like farmers, they might favour strict protection of broodstock in no-take MPAs.</p>
			<p>Some authors have speculated that the implementation of MPAs reduces access to fisheries, resulting in lower catches and revenues for fishermen (e.g. <xref ref-type="bibr" rid="B53">Fletcher et al. 2015</xref> and <xref ref-type="bibr" rid="B25">Chan 2020</xref>). Such studies are sometimes cited as examples of MPAs displacing fishing and negatively impacting resource users. However, the findings of the former have been brought into question by <xref ref-type="bibr" rid="B130">Pecl et al. (2010)</xref> and <xref ref-type="bibr" rid="B82">Hughes et al. (2016</xref>), and the latter has been disproved by research using empirical rather than modelled results, showing that catch and CPUE increased following MPA expansion (<xref ref-type="bibr" rid="B106">Lynham et al. 2020</xref>). <xref ref-type="bibr" rid="B79">Hilborn and Hilborn (2019)</xref> and <xref ref-type="bibr" rid="B117">McConnaughey et al. (2020)</xref> contended that MPAs displace fishing effort, and that this displaced fishing effort then drives down abundance in neighbouring areas, but without evidence of such effects. Indeed, depending on fishery management policies, fishing effort may be displaced, but we found no evidence of any consequent declines in fish abundance or catch outside MPAs. This may be because:</p>
			<list list-type="bullet">
				<list-item>
					<p>the MPA was so small that the fishing effort effect was undetectable,</p>
				</list-item>
				<list-item>
					<p>the fishermen previously active in the MPA discontinued fishing through being compensated (e.g. quotas bought out) and/or changed employment (e.g. to tourism or aquaculture),</p>
				</list-item>
				<list-item>
					<p>there were no data on fishing inside or outside the MPA before or after MPA creation,</p>
				</list-item>
				<list-item>
					<p>some fishing was still allowed in the MPA, as over 94% of MPAs allow fishing (<xref ref-type="bibr" rid="B34">Costello and Ballantine 2015</xref>),</p>
				</list-item>
				<list-item>
					<p>fishing was already ecologically sustainable, negligible or absent in the MPA area prior to establishment (many MPA boundaries are placed to avoid areas important for fishing) or</p>
				</list-item>
				<list-item>
					<p>protection in the MPA counteracted this effect through spillover and larval export.</p>
				</list-item>
			</list>
			<p>It is a credit to how MPAs have been designed and implemented that MPAs have generally benefited fisheries.</p>
			<p>Yet, many MPAs are poorly funded to properly manage and enforce full protection of biodiversity; i.e. too many are &#x201c;paper parks&#x201d; (<xref ref-type="bibr" rid="B135">Relano and Pauly 2023</xref>). Were more MPAs better planned, funded and consequently managed, we would likely see more widespread benefits to biodiversity, including fisheries and people (<xref ref-type="fig" rid="f4">Fig. 4</xref>). The examples in this paper (<xref ref-type="table" rid="t1">Table 1</xref>, <xref ref-type="table" rid="t2">2</xref>) should inspire improved management. The emerging benefits to society will in turn inspire local communities to establish more marine reserves.</p>
			<fig id="f4">
				<label>Fig. 4</label>
				<caption>
					<title>A diagram indicating how protection of marine biodiversity can benefit people and nature.</title>
				</caption>
				<graphic id="gra-4" xlink:href="SCIMAR-88-01-e080-gf4.png"/>
			</fig>
			<p>Several studies estimated negligible fishery losses from the creation of MPAs. For example, the likely costs to fisheries of expanding MPAs in Northern Ireland ranged from &#xa3;0 to &#xa3;6000 per annum per proposed MPA (<xref ref-type="bibr" rid="B41">Department of Agriculture, Environment and Rural Affairs of Northern Ireland 2020</xref>); up to 3.8% loss of income was reported from new MPAs in the Oregon Territorial Sea (<xref ref-type="bibr" rid="B154">The Research Group LLC 2021</xref>); and it was reported that MPA expansion to 30% of the Seychelles EEZ would have a negligible impact on the tuna fishery because the areas only contributed 4% of the catch (<xref ref-type="bibr" rid="B26">Chassot et al. 2018</xref>). If fisheries are already operating in an ecologically sustainable manner, with negligible effect on biodiversity, then creating MPAs may confirm this, and no fishery benefits would be evident, as suggested in a study on the Great Barrier Reef fisheries (<xref ref-type="bibr" rid="B53">Fletcher et al. 2015</xref>).</p>
			<p>Regardless of the reason, not only do claims of fishery displacement effects not seem to have impacted fisheries, but the evidence shows that MPAs sustain or increase catch in adjacent areas (<xref ref-type="table" rid="t1">Table 1</xref>). Rather than negatively affecting fish catch through the displacement of fishing effort, &#x2018;fishing-the-line&#x2019;, where commercial and recreational fishermen concentrate fishing effort along MPA boundaries, has been shown to increase yield and provide greater catches of larger individuals, and is a well-known practice among fishermen (<xref ref-type="bibr" rid="B89">Kelly et al. 2002</xref>, <xref ref-type="bibr" rid="B62">Go&#xf1;i et al. 2006</xref>, <xref ref-type="bibr" rid="B17">Boerder et al. 2017</xref>, personal observation). A metanalysis found fish abundance and biomass increase by 33% and 54%, respectively, immediately outside 23 MPAs (<xref ref-type="bibr" rid="B44">Di Lorenzo et al. 2016</xref>). MPAs increase resilience to fishing, as shown by their ability to sustain consistent catches while subject to intense fishing pressure along their boundaries (<xref ref-type="bibr" rid="B150">Stelzenm&#xfc;ller et al. 2008</xref>, <xref ref-type="bibr" rid="B151">Stobart et al. 2009</xref>, <xref ref-type="bibr" rid="B38">da Silva et al. 2015</xref>, <xref ref-type="bibr" rid="B68">Harasti et al. 2018</xref>). This displacement of fishing effort to reserve boundaries and neighbouring areas may also enhance fish stock availability and stabilize local catches, depending on the relative size of the MPA, as shown for far-ranging pelagic species such as tuna (<xref ref-type="bibr" rid="B17">Boerder et al. 2017</xref>).</p>
			<p>Gear and catch restrictions can reduce impacts on biodiversity, but <xref ref-type="bibr" rid="B76">Hilborn&#x2019;s (2016)</xref> claim that these fishery regulations offer more protection than MPAs is only true when compared with partly protected areas which still allow fishing. Furthermore, most commercial and recreational fishing methods, including hook and line and pots, can kill species of seabirds, mammals, turtles and fish which are already threatened with extinction. Although changes in fishing gear can be successful at reducing bycatch, they do not eliminate it, and technological advances in fishing gear that may reduce bycatch have been slow to be adopted. Furthermore, contrary to assertions by <xref ref-type="bibr" rid="B117">McConnaughey et al. (2020)</xref>, limiting (but not banning) trawl fishing effort cannot have more positive effects on benthic biota than the implementation of MPAs, because bottom trawls destroy biogenic habitats, some created by species with life spans of centuries (<xref ref-type="bibr" rid="B75">Hiddink et al. 2017</xref>). In contrast, MPAs enable the natural recovery of benthic habitats and commercial species (<xref ref-type="bibr" rid="B61">Gell and Roberts 2003</xref>, Stewart et al. 2020).</p>
			<p>The economic benefits of MPAs can be numerous (<xref ref-type="table" rid="t1">Tables 1</xref>, <xref ref-type="table" rid="t2">2</xref>). However, stakeholder negotiation during the design process can lead to MPAs being placed in areas where there is little fishing and low biodiversity, as well as to reductions in protected area size, shape and the level of protection provided (<xref ref-type="bibr" rid="B73">Helson et al. 2010</xref>, <xref ref-type="bibr" rid="B107">Magris and Pressey 2018</xref>, <xref ref-type="bibr" rid="B95">Kuempel et al. 2019</xref>). Sometimes, due to anticipated opposition, MPA boundaries and locations are not designed to maximize benefits to biodiversity and fisheries, but instead are based on political processes that prioritize public acceptance or logistics, ignoring or downgrading ecological and biological aims (<xref ref-type="bibr" rid="B42">Devillers et al. 2015</xref>, <xref ref-type="bibr" rid="B102">Lubchenco and Grorud-Colvert 2015</xref>). This reduces the potential benefits of MPAs, as all outcomes ultimately depend on ecological recovery. MPA design, attributes and stakeholder support play a large part in determining any benefits to fisheries and tourism (<xref ref-type="bibr" rid="B43">Di Franco et al. 2016</xref>), and poorly designed or enforced MPAs may not reap economic benefits (<xref ref-type="bibr" rid="B22">Campbell et al. 2012</xref>). Were more MPAs selected to maximize fishery benefits, economic benefits might be even greater than those found here (<xref ref-type="table" rid="t1">Table 1</xref>). The evidence here strengthens arguments to design both partial-take MPAs and marine reserves to benefit both biodiversity and fisheries rather than shrink them into residual locations. Considering that many, perhaps most, MPAs are not established to benefit fisheries, it is noteworthy that so many show fishery benefits (<xref ref-type="table" rid="t1">Table 1</xref>).</p>
			<p>There do not appear to be any studies demonstrating a clear economic cost to fisheries after the establishment of an MPA, only benefits. <xref ref-type="bibr" rid="B12">Balmford et al. (2004)</xref> estimated that a global MPA network may create 1 million jobs, and its $5-19 billion cost was less than the government subsidies to industrial fisheries, which only serve to postpone the eventual collapse of otherwise unsustainable fisheries and associated employment. They did not provide data on the costs of existing fisheries management, which are likely to be greater than the cost of MPA management (<xref ref-type="bibr" rid="B5">Anonymous 2023</xref>), or estimate the benefits to fisheries. Similarly, a global analysis on the cost-benefits of expanding protected areas on land and sea found that the economic revenue would be $64 to &#xa3;454 billion greater than that of not expanding them by 2050, and would avoid losses of $179 to &#xa3;534 billion (<xref ref-type="bibr" rid="B162">Waldron et al. 2020</xref>). Consequently, the expansion of MPAs, which promote sustainable management, could save money when compared with current fisheries management practices, particularly if the costs of existing fisheries management and subsidies are redistributed. Because of the simplicity of the management, marine reserves have lower management costs than partly protected MPAs and fishery areas (Anonymous 2023). Other studies have also estimated the varying costs of establishing and maintaining protected areas, but similarly do not place this in the context of existing costs in marine spatial management, or consider the costs of continuing the status quo (<xref ref-type="bibr" rid="B84">Jantke et al. 2018</xref>). Estimating the cost-benefits of MPAs to fisheries is complicated, but <xref ref-type="bibr" rid="B19">Brander et al (2020)</xref> conclude that expanding the global MPA network will reap benefits 1.4 to 2.7 times the costs.</p>
			<p>In contrast to land-based agriculture and forestry, fisheries make no investment in habitat or broodstock management. The sea is a public not a private resource, and stakeholders include not only commercial fishermen but also people involved in subsistence and recreational fishing, sport, tourism, education, research, conservation, mining, mariculture and transport. Despite this, the financial costs of MPAs are frequently estimated prior to implementation. Thereafter, the level of protection and the area protected are often reduced, or a financial package is determined, to assuage the temporary loss of resource access to one group of stakeholders (<xref ref-type="bibr" rid="B125">Olsson et al. 2008</xref>, <xref ref-type="bibr" rid="B28">Clifton 2013</xref>). These measures are based on the assumption that wildlife within protected areas belongs to those who exploited it, whereas it is a public resource which the exploiters have not invested in. In fact, protected areas are a method of investing in ecosystem restoration and sustainability, which are the foundations of productive, profitable fisheries and a resource for present and future generations.</p>
			<p>In addition to the lack of research on existing fisheries management costs, the environmental costs of fishing, from carbon emissions to the loss of biodiversity, are seldom found within the literature. In the few studies to estimate environmental costs, the release of greenhouse gases from seabed sediments by bottom trawling is significant (<xref ref-type="bibr" rid="B143">Sala et al. 2021</xref>, <xref ref-type="bibr" rid="B9">Atwood et al. 2023</xref>). Conversely, using protected areas provides a management strategy that benefits biodiversity, which can in turn result in increased carbon capture and storage (<xref ref-type="bibr" rid="B109">Mariani et al. 2020</xref>, <xref ref-type="bibr" rid="B104">Luisetti et al. 2020</xref>, <xref ref-type="bibr" rid="B83">Hutto et al. 2021</xref>, <xref ref-type="bibr" rid="B49">Epstein and Roberts 2022</xref>).</p>
			<p>MPAs can also increase the resilience of biodiversity to climate change through harbouring more abundant and genetically diverse populations (<xref ref-type="bibr" rid="B32">Costello 2021</xref>). For example, studies on abalone in Baja California found that marine reserves enhance resilience to climate impacts in abalone populations, because unfished populations had a larger body size and greater egg production (<xref ref-type="bibr" rid="B119">Micheli et al. 2012</xref>, <xref ref-type="bibr" rid="B120">Mungu&#xed;a-Vega et al. 2015</xref>). More comprehensive economic studies could show additional positive economic benefits of MPAs because of their benefits to fisheries, carbon storage, reducing greenhouse gas emissions, and in some cases, tourism. Further research to address these knowledge gaps is needed.</p>
		</sec>
		<sec id="sec5" sec-type="conclusions">
			<title>Conclusions</title>
			<p>Given recent criticism of MPAs, the challenges faced during their design and designation, and their frequent small size and sub-optimal location, one would expect their economic benefits to be hard to detect or negligible. But the evidence in the scientific literature is that they can provide economic benefits for fisheries and tourism (<xref ref-type="table" rid="t1">Tables 1</xref>, <xref ref-type="table" rid="t2">2</xref>, <xref ref-type="fig" rid="f1 f2 f3">Figs 1-3</xref>). The generality of these benefits across oceans, continents, countries and a diversity of habitats and ecosystems is clear. While such benefits may seem surprising because fishing has been reduced in an area, it is also common sense that unfished stocks will increase in abundance and spread to adjacent areas as adults, juveniles, larvae or eggs. Thus, sweeping dismissals of MPA economic benefits are unfounded.</p>
			<p>Fisheries management already restricts fishing, sometimes with complete bans for years, so it partly already implements no-take MPAs without calling them MPAs. In some areas, widespread fishery controls, such as quota and gear restrictions, already restrict fishing more than MPAs, especially when most MPAs still allow some fishing. An analysis of marine reserves in Sweden found they complemented fishery management measures, but when reopened to fisheries even temporarily the benefits were promptly lost (<xref ref-type="bibr" rid="B16">Bergstr&#xf6;m et al. 2022</xref>).</p>
			<p>MPAs represent a viable, low-tech, cost-effective strategy that can be used effectively for small to large areas (<xref ref-type="bibr" rid="B138">Roberts et al. 2017</xref>). As such, they have proven highly successful, both for safeguarding marine biodiversity and ecosystem functioning, and more pertinently, for reversing fishery declines, securing food provisions and ecosystem services and enabling the sustainable exploitation of fisheries resources (<xref ref-type="bibr" rid="B133">Pitchford et al. 2007</xref>, <xref ref-type="bibr" rid="B88">Jones et al. 2017</xref>, <xref ref-type="bibr" rid="B126">Ortiz-Lozano et al. 2017</xref>) (<xref ref-type="table" rid="t1">Table 1</xref>). Consequently, a review of 118 studies found that no-take, well enforced and older MPAs benefited human well-being (<xref ref-type="bibr" rid="B13">Ban et al. 2019</xref>). MPAs that are accessible to the public and harbour biodiverse habitats and mega-fauna have been shown to generate huge incomes from tourism, providing increased revenue and improved living standards, while contributing significantly to national GDP (<xref ref-type="bibr" rid="B161">Vianna et al. 2012</xref>, <xref ref-type="bibr" rid="B142">Sala et al. 2013</xref>) (<xref ref-type="table" rid="t2">Table 2</xref>). There is also a need to shift the conventional management of fisheries from commercial to include the wider socio-economic benefits to coastal communities (<xref ref-type="bibr" rid="B132">Pitcher and Lam 2015</xref>).</p>
			<p>The literature shows that the largest benefits to fisheries (<xref ref-type="table" rid="t1">Table 1</xref>) and biodiversity come from the designation of marine reserves from which no marine life or materials can be removed (Lester and Halpern 2017, <xref ref-type="bibr" rid="B58">Friedlander et al. 2017</xref>, <xref ref-type="bibr" rid="B141">Sala and Giakoumi 2017</xref>). This &#x201c;Ballantine&#x2019;s Law&#x201d; after the &#x201c;father of marine reserves&#x201d; who championed the then radical idea that MPAs should be completely no-take and permanent following his leading the establishment of the first MPA in New Zealand (which now hosts 44 marine reserves) (<xref ref-type="bibr" rid="B11">Ballantine and Gordon 1979</xref>, <xref ref-type="bibr" rid="B10">Ballantine 2014</xref>, <xref ref-type="bibr" rid="B163">Walls and Gordon 2017</xref>). <xref ref-type="bibr" rid="B34">Costello and Ballantine (2015)</xref> found that 76% of coastal countries had not even one marine reserve, and today they occupy only ~3% of the global ocean (see <ext-link ext-link-type="uri" xlink:href="http://www.mpatlas.org">http://www.mpatlas.org</ext-link> and <ext-link ext-link-type="uri" xlink:href="https://navigatormap.org">https://navigatormap.org</ext-link>).</p>
			<p>The fishing industry and fishing communities have much to gain from MPAs, but misconceptions perpetuated in the scientific literature are serving as barriers to their efficacy and implementation. Global analyses have prioritized where to locate MPAs to meet the calls by the Convention on Biological Diversity, the UN Convention on the Law of the Sea, and the International Union for Conservation of Nature for at least 30% of ocean habitats to be fully protected by 2030 (<xref ref-type="bibr" rid="B169">Zhao et al. 2020</xref>). To achieve this, fishery scientists need to promote the use of MPAs as a strategy to support biodiversity, including &#x201c;ecosystem-based management&#x201d; of fisheries, and work with conservation scientists in order to realize the true capacity of MPAs for economic success (<xref ref-type="bibr" rid="B31">Costello et al. 2016b</xref>, <xref ref-type="bibr" rid="B16">Bergstr&#xf6;m et al. 2022</xref>) (<xref ref-type="fig" rid="f4">Fig. 4</xref>). MPAs are our best strategy for reversing declining biodiversity and unsustainable fisheries, because business as usual for global fisheries is unsustainable.</p>
		</sec>
	</body>
	<back>
		<ack>
			<title>Acknowledgements</title>
			<p>I thank Tamlin Jefferson, John Lynham, Chris McGonigle, Juliano Palacios Abrantes, Joana Smith, Belinda Brambley, Silas Candida Principe De Souza, members of the Octogroup e-mail list community and two anonymous referees and the editor for helpful suggestions that contributed to this paper. Cesc Gord&#xf3;-Vilaseca kindly translated the title and abstract into Spanish.</p>
		</ack>
		<sec sec-type="transparency-statement" id="sec-01">
			<title>Declaration of competing interest</title>
				<p>The author of this article declares that he has no financial, professional or personal conflicts of interest that could has inappropriately influenced this work.</p>
		</sec>
		<sec sec-type="apoyo" id="sec-02">
			<title>Funding sources</title>
				<p>This paper is a contribution to the project MPA Europe funded by Horizon Europe under Grant Agreement 101059988.</p>
		</sec>
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