Designate a Marine Protected Area and prohibit all types of fishing
Overall effectiveness category Likely to be beneficial
Number of studies: 29
Background information and definitions
Fishing can impact subtidal benthic invertebrates through species removal or habitat damage from fishing gear entering in contact with the seabed (Collie et al. 2000). Specific areas can be designated as protected, and specific management measures taken to control for impactful activities, such as prohibiting all types of fishing (Villamor & Becerro 2012). Such areas are often referred to as marine reserves, marine sanctuaries, or more commonly “no-take areas/zones”. Inside such protected no-take areas, the threat to subtidal benthic invertebrates is removed, and previously impacted populations are, in theory, able to recover over time (Jack & Wing 2010).
When this intervention occurs outside of a protected area, evidence has been summarised under “Threat: Biological resource use – Cease or prohibit all types of fishing”.
Collie J.S., Hall S.J., Kaiser M.J. & Poiner I.R. (2000) A quantitative analysis of fishing impacts on shelf‐sea benthos. Journal of Animal Ecology, 69, 785–798.
Jack L. & Wing S.R. (2010) Maintenance of old-growth size structure and fecundity of the red rock lobster Jasus edwardsii among marine protected areas in Fiordland, New Zealand. Marine Ecology Progress Series, 404, 161-172.
Villamor A. & Becerro M.A. (2012) Species, trophic, and functional diversity in marine protected and non-protected areas. Journal of Sea Research, 73, 109–116.
Supporting evidence from individual studies
A site comparison study in 1988–1994 in two sandy areas with seagrass in Exuma Cays, North Atlantic Ocean, central Bahamas (Stoner & Ray 1997) found that inside a protected marine reserve closed to all fishing for 33–36 years, abundance of adult and larval queen conch Strombus gigas were higher, but abundance of juveniles was similar, compared to a nearby unprotected fished area. Adult conch abundance was higher in the closed area (34–270 conch/ha), compared to the fished area (2–88 conch/ha). In six of 13 comparisons, larval abundance was significantly higher in the closed area (1–50 larvae/10 m3) compared to the fished area (0.06–6), and statistically similar in seven comparisons (closed: 0.2–55; fished: 0.25–1.6 larvae/10 m3). Juvenile conch abundance was statistically similar in closed (2–6 conch/ha) and fished areas (0–2). The marine reserve (456 km2) was designated in 1958, closed to all fishing and the collection of any animals prohibited. Inside the fished area, taking juvenile queen conch and using SCUBA gear for fishing is prohibited. A snorkeller counted adult queen conch along 12 transects (6 m wide; total area of 28 ha) in March–September 1991 (fished area) and 1994 (closed area). Divers also measured adult conch abundance, shell length and lip thickness (see paper for details). Juvenile conch abundance was estimated in each area (annually in 1988–1991 in the fished area; in 1991 in the closed area; see paper for details). Queen conch larvae were surveyed in June–August 1993–1994 using plankton nets.Study and other actions tested
A paired, site comparison study in 1992–1997 of six coralline seabed areas in the South Pacific Ocean, northeastern New Zealand (Cole & Keuskamp 1998) found that inside marine protected areas prohibiting all fishing for more than 10 years Kina sea urchins Evechinus chloroticus tended to be heavier compared to adjacent fished areas, but that all areas appeared to have similar urchin abundance. Results were not statistically tested. Sea urchins tended to be heavier inside protected areas (average 54 g), compared to fished areas (40 g). Sea urchin abundance varied between 1 to 4 urchin/m2 in protected areas, and 1 to 7 urchin/m2 in fished areas. Three marine protected areas prohibiting all exploitation (assumed to include all fishing) and three paired adjacent fished areas were sampled between 1992 and 1997 (4–5 sites/area; 5–10 m depth). Divers counted all urchins in twenty 1 m2 quadrats and measured the diameter of at least 60 urchins. In one protected and one fished area, the skeletons of ten sea urchins (55–65 mm diameter)/site were wet-weighed.Study and other actions tested
A replicated, paired, site comparison study in 1995 in eight sites of kelp beds around islands in the South Pacific Ocean, northeastern New Zealand (Babcock et al. 1999) found that marine protected areas prohibiting all fishing (no-take) for 20 years had more and bigger spiny lobsters Jasus edwardsii compared to adjacent fished areas. Abundance of lobsters was greater in no-take (455 lobsters/ha) compared to fished areas (174 lobsters/ha). In addition, average lobster size was greater in no-take (110 mm), compared to fished areas (94 mm). Leigh Marine Reserve (established 1975) was surveyed in spring 1995, and Tāwharanui Marine Park (established 1982) in autumn 1995. At two sites within and two sites outside each no-take area, divers counted and visually estimated the carapace length of all lobsters within five randomly-placed 50 × 10 m transects (max. 25 m depth).Study and other actions tested
A replicated, site comparison study in 1998 of eight rocky and sandy areas in the San Juan Archipelago, northwest Pacific Ocean, USA (Tuya et al. 2000) found that the effects of prohibiting all fishing and harvesting within marine protected areas (no-take) on the abundance of red sea urchins Strongylocentrotus franciscanus depended on the age of the protected area and the size of the urchins, and that no-take areas did not affect abundances of sea cucumbers Parastichopus californicus and scallops Chlamys rubida, Chlamys behringiana, and Hinnites giganteus. Abundances of medium and large urchins were higher in the eight-year-old no-take areas (medium: 65; large: 225 individuals/300 m2) compared to one-year-old no-take areas (medium: 21; large: 43) and unprotected areas (medium: 9; large: 21). There were no significant differences between young protected areas and unprotected areas. Abundance of small urchins was similar across areas (1–4). Abundance data for sea cucumbers and scallops were not provided. The authors suggest the lack of increase in abundances inside protected areas was likely due to a lack of compliance and enforcement of prohibitions. In July 1998, three marine preserves (established eight years prior and prohibiting the harvest of organisms; sea urchin fishery closed since the late 1970s), two marine protected area (designated in 1997; voluntary no-take zones), and three unprotected areas were surveyed. Divers counted and measured red sea urchins, sea cucumbers, and scallops along 300 m2 transects (4 transects/area).Study and other actions tested
A site comparison study in 1994–1995 in four coral reef areas in Exuma Sound, North Atlantic Ocean, Bahamas (Lipcius et al. 2001) found that the effects of prohibiting all fishing within marine protected areas (no-take) on the abundance and size of Caribbean spiny lobsters Panulirus argus after 8–9 years varied between areas, year and seasons. Overall, lobster abundance was higher in the no-take area (63–51 lobsters/ha) than in only one of three fished areas (data not shown), and similar to abundance in the other two fished areas (6–333 lobsters/ha). The closed area had larger males (118 mm) than two (105–106 mm) of three fished areas (no difference to the other; 113 mm). However, the closed area had larger females (109 mm) than only one (95 mm) of three fished areas (no difference to the other two; 108–128 mm). In 1986, a 456 km2 marine reserve (established in 1959) was designated as a no-take area (although some poaching still occurred). In spring (before the fishing season) and autumn (after the fishing season) 1994–1995, lobster abundance was recorded in the reserve and in three adjacent fished areas (3–17 reef sites/area). Divers examined crevices at depths <20 m, and measured (carapace length), and sexed all lobsters found.Study and other actions tested
A site comparison study in 1994–2000 of twelve rocky seabed and soft sediment sites in Tonga Island Marine Reserve, Tasman Sea, New Zealand (Davidson et al. 2002) found that five to seven years after designating a marine protected area prohibiting all fishing (no-take reserve), abundance of spiny lobsters Jasus edwardsii was similar inside the protected area, compared to outside where fishing occurred, but lobster size was higher. Average abundance was not statistically higher inside the reserve (1.3 lobsters/100 m2) compared to outside (0.4 lobsters/100 m2). Lobster were larger inside the reserve 113–132 mm), compared to outside (94–104 mm; results not statistically tested). In addition, when compared with data from 1994 (12 months after establishing the reserve) average lobster abundance increased inside the reserve (1994: 1; 1998–2000: 1.4 lobster/100 m2) but decreased outside (1994: 0.7; 1998–2000: 0.6 lobster/100 m2; results not statistically tested). Tonga Island Marine Reserve was established in 1993 and is closed to all fishing and harvesting. Seven sites inside and five sites outside the reserve (6–11 m depth) were surveyed on six occasions in 1998–2000. During each survey, divers counted and estimated the size of lobsters in twelve 24 x 4 m transects/site. Abundance data from 1998 and 2000 (obtained in December) were compared with a prior survey in December 1994 of five sites inside and five sites outside the reserve (10–12 transects/site; 30 × 4 m).Study and other actions tested
A site comparison study in 1983–2001 of three sites of kelp forest in the Channel Islands National Park, southern California, North Pacific Ocean, USA (Rogers-Bennett et al. 2002) found that the effects of designating protected areas prohibiting all fishing (no-take) on abundance, size, and egg production of pink abalone Haliotis corrugata depended on the level of enforcement. Five to 23 years after their designation, cumulative abundance across years was higher in an enforced (333 abalone/13,040 m) compared to a not enforced (116) no-take area, but these were not significantly different from a fished site (431). Abundance declined over time at all sites. Size of abalones was higher in the enforced no-take area (147 mm) and in the unenforced no-take area (134 mm) compared to the fished site (122 mm). More large abalone (above minimum landing size of 158 mm) were found in the enforced no-take area (30%) and in the unenforced no-take area (6%) compared to the fished site (2%). Egg production was higher in the enforced no-take area (2,555; units unclear) compared to the other sites (unenforced no-take: 550; fished site: 1,420). Annually between 1983 and 2001, pink abalone were counted and measured by divers along 10–12 transects (40–60 m2) at three sites. Two were no-take areas established in 1978 (one enforced, one not enforced) and one a site where non-commercial fishing occurred (commercial fishing was prohibited). Egg production was estimated from abundance and size data.Study and other actions tested
A site comparison study in 1999–2001 of eight sites in two rocky areas in the South Pacific Ocean, New Zealand (Shears & Babcock 2003) found that designating a marine protected area prohibiting all fishing (no-take reserve) had mixed effects on invertebrate abundances depending on species, after 23–25 years. Abundance of Cook’s turban snails Cookia sulcata was higher at sites inside the reserve (2–16 individuals/m2) compared to fished sites outside (1–2), but abundance of star limpets Cellana stellifera was lower inside (0–2). compared to outside (0–6). Abundances were similar at sites inside and outside the reserve for the green top shells Trochus viridis (inside: 0–12; outside: 0–10) and red opal top shells Cantharidus purpureus (inside: 1–14; outside: 0–22). Leigh Marine Reserve (no-take area) was established in 1975 (date taken from (3) summarised above). Annually in summer in 1999–2001, four sites inside and four outside the reserve were sampled. Invertebrates were counted in twenty 1 m2 quadrats/site (2–10 m depth).Study and other actions tested
A replicated, randomized, site comparison study in 1997–2001 in two areas of rocky reef in the Florida Keys, USA (Cox & Hunt 2005) found that inside a protected marine reserve prohibiting all fishing and harvesting, the size and abundance of Caribbean spiny lobsters Panulirus argus were higher compared to outside the reserve where seasonal fishing was allowed, but the population sex ratio was similar. On average over the four years following its designation, lobsters were bigger inside the reserve (82–94 mm carapace length), compared to outside (77–85 mm). There were more lobsters bigger than 76.2 mm (legal-catch size) in the reserve (3–86/60 min search) compared to outside (0.7–71/60 min search). The population sex ratio was not significantly different inside (0.5–3 female:male) and outside the reserve (0.4–5). In addition, inside the reserve, average lobster sizes increased during the study (1997: 82–86; 2001: 87–94 mm). In July 1997, a 3,000-ha reserve was established prohibiting all fishing and harvesting year-round. Annually in July and September of 1997–2001, divers counted, measured, and recorded the sex of lobsters during twenty-four 60 min timed-surveys at randomly-chosen locations; twelve inside and twelve outside the reserve.Study and other actions tested
A replicated, site comparison study in 2002–2003 in one area of mixed seabed off the coast of southern California, North Pacific Ocean, USA (Parnell et al. 205) found that a protected marine reserve prohibiting all fishing had higher abundances of three out of five surveyed invertebrate species compared to outside after approximately 30 years. Average abundances were higher inside the reserve than outside the reserve for red sea urchins Strongylocentrotus franciscanus (inside: 0.86/m2 vs outside: 0.02/m2), rock scallops Crassadoma giganteum (0.006/m2 vs 0.001/m2), and green abalone Haliotis fulgens (0.007/m2 vs 0.001/m2), but not for California spiny lobsters Panulirus interruptus (0.006 vs 0.004/m2) and pink abalone Haliotis corrugata (0.003/m2 vs 0.004/m2). A 2.16 km2 area was established as a protected no-take marine reserve in 1971. Between spring and summer 2002, a total of 286 transects (30 × 4 m) were surveyed by divers inside and outside of the reserve (numbers of transects unspecified), and the abundances of red sea urchins, rock scallops, spiny lobsters, and pink abalone recorded. Green abalone abundance was recorded during dive surveys along 500 m transects (numbers unspecified) at depths below 6 m between spring and autumn 2003.Study and other actions tested
A replicated, paired, site comparison study in 2002–2003 of six areas of muddy seabed and seagrass in Moreton Bay, Coral Sea, Australia (Pillans et al. 2005) found that marine protected areas prohibiting all fishing (no-take reserves) had typically more and bigger mud crabs Scylla serrata, compared to nearby and distant fished areas, five years after designation. Abundance of mud crabs was approximately 2.5 times higher in the reserves (1.5 crabs/pot) compared to three of four fished areas (0.3–0.5 crabs/pot), but not significantly different from one nearby fished area (1.1 crabs/pot). Mud crabs were on average bigger in the reserves (15.7–16.1 cm), compared to the fished areas (14.7–15.8 cm). Two no-take marine reserves (closed to all fishing) were established in 1997. Mud crabs were surveyed in the reserves and in four fished (recreationally and/or commercially, see paper for details) non-reserves (two paired with each reserve; one nearby ≤7 km away, one distant (distance unspecified)) in summer and winter 2002–2003. During each survey, 11 baited crab pots/area were deployed at 1–4 m depth (≥50 m apart), for 24 h and recovered (repeated two consecutive days). Upon recovery, all crabs captured were identified, measured (carapace width), and released. The relative abundance of crabs was expressed as catch/unit effort (meaning the number of crabs caught/pot).Study and other actions tested
A replicated, before-and-after, site comparison study in 1977–2005 of 10 rocky sites in the South Pacific Ocean, northeastern New Zealand (Shears et al. 2006) found that during the 22 years after implementing a protected marine park prohibiting all fishing (no-take), abundance and biomass of spiny rock lobsters Jasus edwardsii increased and became greater than at adjacent fished sites. Before designation, lobster abundance and biomass were similar inside (4–19 lobsters/transect; 0–3 kg/transect) and outside (0–7 lobsters/transect; 0.1 kg/transect) the park. Legal-size lobsters (>95 mm carapace length) in the park were 10.9 times more abundant after implementation (6–32 lobsters/transect) compared to before (0–5 lobsters/transect), with biomass 25 times higher (before: 0–3; after: 14–41 kg/transect). There was no change in abundance of sublegal-size lobsters inside the park. No legal-size and only 0–7 sublegal-size lobsters/transect were present outside the park after implementation. Tāwharanui Marine Park was established in 1981 (implemented 1983). Between 1977 and 2005, five no-take sites and five fully-fished sites outside the park were surveyed annually. Divers counted all lobsters and visually estimated the size and weight of legal-size lobsters along a 50 × 10 m fixed transect at each site.Study and other actions tested
A review of 14 studies undertaken between 1985 and 2002 in 20 areas of seabed in New Zealand (Pande et al. 2008) found that marine reserves prohibiting all fishing (no-take) typically had bigger and more abundant spiny rock lobsters Jasus edwardsii compared to fished areas outside the reserves. In 12 of 13 studies, rock lobsters were bigger inside the reserves (98 mm) than outside (79 mm), and in 11 of 14 studies lobster were more abundant inside the reserves (0.03 lobsters/m2) than outside (0.01 lobsters/m2). Older and larger reserves had greater effects than younger and smaller ones on lobster size (data presented as effect sizes). Size and abundance data were extracted from 14 studies of 10 marine reserves and 10 corresponding fished areas and used in a meta-analysis. At the time of surveys, the reserves were on average 8.5 years old.Study and other actions tested
A before-and-after, site comparison study in 1992–2007 in twelves rocky sites in the Tasman Sea, Australia (Alexander et al. 2009) found that over the 15 years after designating and enforcing a marine protected area prohibiting all fishing, mobile macroinvertebrate species richness remained stable at protected sites but decreased at fished sites, while overall abundance did not change at any sites. Before enforcement, total macroinvertebrate species richness was lower at protected sites (11 species) compared to fished sites (16). After 15 years, species richness remained stable within protected sites (10–12) but had decreased in fished sites to similar levels (13–14). Before enforcement, overall mobile macroinvertebrate abundance was lower at protected sites (330–560 individuals/site) than fished sites (760–1,030) and remained similar at all sites over 15 years (protected: 375–430; fished: 625–820). This pattern was due to opposing changes in abundances of specific groups and species (see paper for details). In addition, abundance of blacklip abalone Haliotis rubra decreased over time inside the protected sites relative to fished sites, while abundance of southern rock lobsters Jasus edwardsii increased in protected sites but decreased in fished sites (data not provided). An area within Maria Island National Park was declared a no-take area in 1991 and closed to all fishing. In spring 2006 and autumn 2007, a diver visually identified and counted all mobile macroinvertebrates (echinoderms, crustaceans, and molluscs >1 cm) along four 50 m transects at six sites inside and six outside the no-take area (5 m water depth). Data were compared to historical surveys in spring and autumn 1992 before effective enforcement.Study and other actions tested
A replicated, site comparison study in 2006–2007 of 11 rocky seabed sites in Fiordland, Tasman Sea, New Zealand (Jack et al. 2009) found that the effects of marine protected areas prohibiting all fishing (no-take reserves) on the abundance of red rock lobsters Jasus edwardssi and percentage cover of its prey, the mussels Mytilus edulis galloprovincialis, Perna canaliculus, and Aulacomya maoriana, compared to a commercial fishing exclusion zone, varied with the age and location of the protected areas. Lobster abundance was higher in the >13-year-old no-take reserve established in 1993 (36 individuals/250 m2) compared to one of two <2-year-old no-take reserves established in 2005 (Kutu Parera; 11 individuals/250 m2) and the commercial exclusion zone (9 individuals/250 m2). The second <2-year-old no-take reserve (Taipari Roa) had no lobsters. Mussel cover was higher in the >13-year-old reserve (28%) compared to the exclusion zone (15%), and cover at Kutu Parera (18%) was not different from either the >13-year-old reserve or the exclusion zone. No mussels were found at Taipari Roa. Lobsters and mussels were surveyed by divers at two sites in each of the following: a no-take reserve established in 1993, and two no-take reserves established in 2005, and at five sites within a commercial exclusion zone set in 2005 (15 m depth). During six surveys in 2006–2007, red rock lobsters were counted along 50 × 10 m transects (1–4 transects/site/survey). The percentage cover of mussels (species combined) was estimated from 25 photographs (0.17 m2)/site taken during a single survey in 2007.Study and other actions tested
A systematic review of 149 studies published between 1977 and 2006 of no-take marine reserves across the world (Lester et al. 2009) found that inside marine protected areas prohibiting all fishing, invertebrate biomass, abundance, and size were greater, but species richness was not, compared to unprotected areas outside. Inside the reserves, average biomass increased by 752%, average abundance by 176%, and average size by 26%, compared to outside the reserve. Species richness decreased by a non-significant <5% inside compared to outside the reserves. When analysed by species group, molluscs and arthropods had the greatest increases (molluscs: +240% (non-significant) biomass, +422% abundance, +33% size; arthropods: +889% biomass, +323% abundance, +33% size), while there were no significant changes for any metrics for echinoderms or cnidaria. Species highly targeted by fisheries had the greatest increases in abundance (+385%) and biomass (+820%) in the reserves. The selected studies compared invertebrate abundance, biomass, size, and species richness for 124 reserves across 29 countries. Selected studies included comparisons of before-and-after the reserves were established, and comparisons of inside vs outside the reserves. A meta-analysis was performed on the selected studies.Study and other actions tested
A replicated, site comparison study in 2006–2007 of 26 rocky seabed sites in Fiordland, Tasman Sea, New Zealand (Jack & Wing 2010a) found that older marine protected areas prohibiting all fishing (no-take reserves) had more red rock lobsters Jasus edwardsii compared to younger ones, to a commercial fishing exclusion zone and to adjacent areas without designated protection. Lobster abundance was higher in no-take reserves >13-year-old established in 1993 (12 individuals/250 m2) compared to those <2-year-old established in 2005, commercial exclusion zones, and adjacent unprotected areas which had similar abundances to each other (1–2 individuals/250 m2). In 2006 and 2007, divers surveyed four no-take reserves established in 1993, ten no-take reserves established in 2005, eight sites within a commercial fishing exclusion zone set in 2005, and four unprotected fished sites. All sites were located at 15 m depth on rocky habitat. Red rock lobsters were counted along 50 × 5 m transects (1 transect/site in 2006, 3/site in 2007).
A replicated, site comparison study in 2008 of eight rocky seabed sites in Fiordland, Tasman Sea, New Zealand (Jack & Wing 2010b) found that marine protected areas prohibiting all fishing (no-take reserves) had different population structures of female red rock lobsters Jasus edwardsii, but not males, and greater egg production potential, compared to commercial fishing exclusion zones, but the effects varied with the age of the reserves. Population structure data were reported as size-frequency distributions. A 15-year-old reserve had greater abundance and size of female lobsters compared to commercial exclusion zones. One of two 3-year-old reserves had no lobsters (either male or female); in the other abundance and size of female lobsters were not significantly different to the other sites. Egg production was higher in the 15-year-old reserve (8,350/m2/year) compared to the commercial exclusion zones (1,260/m2/year). The 3-year-old reserve with lobsters had an egg production not significantly different to the other sites (3,400/m2/year). In 2008, divers surveyed two sites in each of the following: a no-take reserve established in 1993, two no-take reserves established in 2005, and a commercial fishing exclusion zone set in 2005. All sites were located at 15 m depth on rocky habitat. Red rock lobsters were counted along three 50 × 5 m transects, and their size and sex assessed from video footage (see study for details). Egg production potential was estimated using abundance and size data for female lobsters.Study and other actions tested
A before-and-after, site comparison study in 1997–2009 in areas of soft seabed in the western Mediterranean Sea, off the coast of Sardinia, Italy (Follesa et al. 2011) found that, over the 10 years after designating a marine protected area prohibiting all fishing (no-take), abundance and biomass of European spiny lobster Palinurus elephas increased inside the no-take area, and after 10 years was greater than in an adjacent fished area. Within the no-take area lobster abundance increased over time (1998: 0.4 lobsters/net; 2009: 1.5 lobsters/net) and so did biomass (1997: 0.09 kg/net; 2009: 0.56 kg/net). In 2008–2009 (after 10 years) lobster abundance within the no-take area was 4.7 times greater than in the fished area (data not shown). The Su Pallosu no-take area was closed to fishing in 1998. Annually from 1997 to 2009, trammel nets (50 m long) were towed inside the no-take zone (91 nets in total; 2–14 nets/year). Similar nets were towed in an adjacent fished area in 2008 and 2009 (256 nets; within 25 km2 of the no-take zone). Lobsters in each net were counted and weighed, and total abundance and biomass for each area estimated.Study and other actions tested
Referenced paperFollesa M.C., Cannas R., Cau A., Cuccu D., Gastoni A., Ortu A., Pedoni C. & Porcu C. (2011) Spillover effects of a Mediterranean marine protected area on the European spiny lobster Palinurus elephas (Fabricius, 1787). Aquatic Conservation: Marine and Freshwater Ecosystems, 21, 564-572.
A replicated, site comparison study in summer 2004–2007 of six sites in three rocky and sandy seabed areas in the Bristol Channel and the Irish Sea, UK (Hoskin et al. 2011) found that a marine protected area prohibiting all fishing (no-take) had mixed effects on the abundances and sizes of European lobster Homarus gammarus, velvet crab Necora puber, brown crab Cancer pagurus and spider crab Maja squinado. Abundance of large lobsters (≥90 mm) increased by 127% inside the no-take zone between 2004 and 2007 (from 3 to 7 lobsters/line) and was five times higher than in unprotected fished areas where abundance had not changed (1–2 lobsters/line). Abundance of small lobsters (<90 mm) increased by 97% (from 3 to 7 lobsters/line) in the no-take zone, but remained constant in the fished areas (2–4 lobsters/line). The size of large lobsters (≥90 mm) increased by 5% inside the no-take zone between 2004 (98 mm) and 2007 (103 mm) and became 9% larger than in the fished areas where lobster size decreased by 2% (from 98 to 95 mm). The size of small lobsters did not change over time and was similar across all areas. Abundance of velvet crabs decreased by 65% inside the no-take zone over time (from 2 to 1 crabs/line; likely due to increased predation by lobsters) but increased in the fished areas (from 0–6 to 1–7 crabs/line). The average size of velvet crabs did not change over time and was similar across all areas. Abundance of brown crabs did not change over time inside the no-take zone (0.3 crab/line), nor in the fished areas (from 0.3–2 crabs/line). The average size of brown crabs increased by 25% inside the no-take zone between 2004 (115 mm) and 2007 (144 mm) but was not greater than in fished areas (116–130 mm). Abundance of spider crabs was similar in 2004 and 2007 for all areas but varied spatially (with the no-take zone having a lower abundance). The average size of spider crabs did not change over time inside the no-take zone. Lundy Island Marine Protected Area was designated as a voluntary reserve in 1971 (statutory since 1986). In 2003, it included a 4 km2 no-take zone (no fishing or harvesting allowed), the rest being a refuge zone only allowing crab and lobster potting. In 2004–2007, lobsters and crabs were surveyed inside the no-take zone and two unprotected fished locations (20–100 km away) (2 sites/location). Four lines of standard commercial baited shellfish pots were deployed (10 pots/line) at each site for 24 h. Upon retrieval, lobsters and crabs were counted and measured (carapace length). The pots were redeployed for five consecutive days each year.Study and other actions tested
A replicated, site comparison study in 2004 in two rocky reef areas in the Tasman Sea, Australia (Ling & Johnson 2012) found that marine protected areas prohibiting all fishing (no-take reserves) had more spiny rock lobsters Jasus edwardsii compared to fished areas outside the reserves. After 12 and 33 years, average lobster abundances were higher inside the two reserves (12 years: 0.8; 33 years: 1.7 lobsters/100 m2), compared to outside where lobsters were absent (0 lobster/100 m2). Maria Island Marine Reserve was established in 1992. Crayfish Point Marine Reserve was established in 1971. Abundance of lobsters bigger than 140 mm carapace length was recorded along six 50 × 4 m transects during 15 min standardized timed-searches (six inside and six outside each reserve).Study and other actions tested
A replicated, paired, site comparison study in 2008 of ten rocky seabed areas in the Mediterranean Sea, Spain (Villamor & Becerro 2012) found that marine protected areas prohibiting all fishing (no-take) for at least 10 years had similar overall combined invertebrate, fish, and algae community composition and species diversity to unprotected fished areas. Overall community composition varied geographically, but not with protection status (data presented as graphical analyses). Species diversity was similar across areas (diversity presented as a diversity index). In addition, two of ten filter-feeding invertebrate groups were more abundant inside the no-take areas than outside (see paper for details). In August 2008, three sites inside each of five no-take areas (designated between 1983 and 1998) and three sites in each of five adjacent (paired) fished areas were surveyed at 4–11 m depth. All organisms (invertebrates, fish, and algae), were identified and counted along a 50 m2 transect/site. Filter-feeding invertebrates were sorted into 10 trophic groups.Study and other actions tested
A replicated, site comparison study in 2006–2007 of twelve rocky reef sites in the Tasman Sea, Australia (Alexander 2013) found that sites within a marine reserve prohibiting all fishing for 16 years had statistically similar combined mollusc and echinoderm species richness, abundance, and community composition to sites outside the reserve subject to fishing. Species richness and abundance data were not reported. Community composition data were reported as a graphical analysis. Abundance data were reported for the commercially valuable but declining blacklip abalone Haliotis rubra and was lower inside (0.25–0.29 abalone/boulder) compared to outside (0.62–0.94;) the reserve. In summer 2006–2007, ten boulders (30 × 30 × 5 cm) were deployed at each of 12 sites: six within the reserve (declared in 1991) and six outside. Boulders were recovered in January 2007 after three months, and molluscs (including blacklip abalone) and echinoderms present on their underside identified and counted. This was repeated in autumn and winter 2007.Study and other actions tested
A before-and-after, site comparison study in 2004 and 2012 in 18–19 sites of sandy and coral seabed in Tatakoto Atoll, South Pacific Ocean, French Polynesia (Andréfouët et al. 2013) found that over the eight years after designating a marine protected area prohibiting all fishing (no-take), abundance and biomass of giant clams Tridacna maxima appeared higher inside compared to outside the no-take area, but were decreasing in both areas. Results were not tested for statistical significance. Abundance of clams decreased inside (from 119 to 11 clams/m2) and outside (from 9 to 3 clams/m2) the no-take zone. Abundances in 2012 corresponded to only 9.5% of the 2004 abundance for inside and 41% for outside the no-take zone. Total biomass of clams also decreased inside (from 256 to 32 tonnes) and outside (from 126 to 61 tonnes) the no-take zone. Authors state that decreases were linked with mass-mortalities occurring in 2009 due to a high range of temperature variations. In 2004 a 0.5 km2 no-take zone was designated for giant clam conservation in Tatakoto Atoll; the rest of the Atoll is open to hand-harvest of clams by local fishers only. Five to six sites inside the no-take zone and 13 sites outside (within the Atoll) were surveyed in 2004 before designation and in 2012. Snorkelers counted and measured all clams inside six 0.25 m2 quadrats/site (methods fully described in Gilbert et al. 2006).
Gilbert A., Andréfouët S., Yan L. & Remoissenet G. (2006). The giant clam Tridacna maxima communities of three French Polynesia islands: comparison of their population sizes and structures at early stages of their exploitation. ICES Journal of Marine Science, 63, 1573–1589.Study and other actions tested
A systematic review of 14 studies published until February 2011 of marine protected areas across the world prohibiting all types of fishing (no-take) (Sciberras et al. 2013) found that they had more lobsters (species unspecified) compared to marine protected areas only partially prohibiting fishing. Lobster abundance was on average 1.76 times higher in no-take areas compared to partially protected areas (data were not reported, but the analysis outcome was reported as statistical model results). The selected studies compared lobster abundance inside 14 no-take areas with adjacent partial fishing prohibition zones. The abundance data were extracted and used in a meta-analysis.Study and other actions tested
A paired, site comparison study in 2010–2013 in one soft sediment area in the Firth of Clyde, west coast of Scotland, UK (Howarth et al. 2015) found that five years after designation, a marine reserve closed to all fishing had similar species richness, diversity and overall community composition of large invertebrates compared to an adjacent fished area, but higher species richness, similar diversity, and different overall community composition of small sessile invertebrates. Diversity was reported as diversity indices, and community composition as a graphical analysis. After five years, the closed area had similar total abundance of large invertebrates (60/100 m2) compared to the fished area (45/100 m2). However, the closed area had higher abundance of feather stars (closed: 18 vs fished: 11 individuals/100 m2) and eyelash worms Myxicola infundibulum (3 vs 1 individuals/100 m2). Abundances were similar inside and outside the closed area for large crustaceans and starfish combined (32 vs 29 individuals/100 m2), and for the parchment worms Chaetopterus spp. (6 vs 4 individuals/100 m2). After five years, species richness of small sessile invertebrates was higher in the closed area (5 species/m2) than the fished area (4 species/m2). In addition, abundances (as % cover) were higher inside than outside the protected area for hydroids (inside: 4% vs outside: 3%) and sponges (0.3% vs 0.1%), but abundances were similar inside and outside for bryozoans (0.5%) and for worms, anemones, and tunicates (data not provided). Lamlash Bay marine reserve (2.67 km2) was established in September 2008 and closed to all fishing. Annually between 2010 and 2013 at 0–30 m depth, 14–20 sites were samples inside and outside the reserve. Large (size unspecified) invertebrates were counted inside 150 m2 transects. Forty 1 m2 quadrats/transect were photographed and any small sessile organism (invertebrates including corals and algae) present were identified.Study and other actions tested
A systematic review of 150 studies published between 1977 and 2012 of marine protected areas prohibiting all fishing (no-take reserves) across the world (Huijbers et al. 2015) found that they had more invertebrates compared to fished areas outside. Abundance data were not reported, but the analysis outcome was reported as statistical model results. However, the effect of the reserves on invertebrate abundances differed by trophic group. Abundances were greater inside the reserves compared to outside the reserves for omnivorous and filter-feeding invertebrates, but not for carnivorous and herbivorous invertebrates. The proximity of the reserves to coastal influences (landmasses, urban centres, rivers, populations) did not affect the positive effect of marine reserves on invertebrate abundance. The selected studies compared invertebrate abundances inside and outside 113 no-take reserves. A total of 1,416 abundance comparisons were used in a meta-analysis.Study and other actions tested
A replicated, randomized, site comparison study in 2000–2014 in two areas of mixed seabed in the northwestern Mediterranean Sea, Spain (Díaz et al. 2016) found that inside a marine protected are prohibiting all fishing (no-take reserve for lobsters) biomass, abundance, and egg production of Caribbean spiny lobsters Palinurus elephas were higher, and population size structures were different, compared to a fished area outside the reserve. After prohibiting fishing for 21 to 25 years, lobster biomass and abundance had increased inside the reserve, and were higher compared to outside (data reported as indices). Maximum sizes of lobsters were higher inside the reserve (female: 172; male: 190 mm) compared to outside (female: 130; male: 140). Egg production was higher inside the reserve (3.5 million eggs/unit area) compared to outside (84,000–137,000 eggs/unit area). In 1990, a 55 km2 marine reserve was established prohibiting all lobster fishing. Lobster catch data between 2000 and 2014 were obtained from fishing nets deployed inside (total number of nets: 252) and at locations outside the reserve (1–150 km away; total number of nets: 2,671).Study and other actions tested
A replicated, site comparison study in 2013 of twenty sites in the Paterson Inlet, Foveaux Straight, New Zealand (Twist et al. 2016) found that sites within a marine protected area prohibiting all fishing (no-take reserve) had more New Zealand scallops Pecten novaezealandiae after nine years compared to adjacent sites in a recreational harvest-only area, but fewer than at sites in a customary fisheries area. Scallop abundance was higher inside the no-take reserve (0.63 scallops/m2) compared to the recreational area (0.56 scallops/m2), but not compare to the customary area (3.62 scallops/m2). Scallop measured on average 110 mm inside the no-take reserve, 132 mm in the recreational area, and 104 in the customary fisheries area (differences not statistically tested). In June 2013, divers counted and measured scallops in three to nine transects (100 m2) at each of 20 sites: three in the no-take reserve (designated in 2004), three in the recreational harvest-only area, and six in the customary fisheries area (community-based management, see paper for details).Study and other actions tested
A site comparison study in 2012–2015 in eight rocky seabed areas in the Firth of Clyde, west coast of Scotland, UK (Howarth et al. 2017) found that the effects of prohibiting all fishing in a marine reserve on the abundances and sizes of European lobsters Homarus gammarus, brown crabs Cancer pagurus and velvet swimming crabs Necora puber, compared to fished areas, varied between the species. After seven years (in 2015) inside the closed area, there were more European lobsters (1 lobster/pot) and velvet swimming crabs (1.3 crab/pot), compared to fished areas (lobsters: 0.4–0.9/pot; crabs: 0.6–0.7/pot), but fewer brown crabs (closed: 0.8/pot; fished: 1.1–1.3/pot). Over the four-year study, the closed area had on average larger lobsters (closed: 100 vs fished: 82–89 mm) and velvet swimming crabs (71 vs 69 mm) compared to the fished area, but smaller brown crabs (125 vs 147 mm). In addition, over the study period, lobster sex ratios (male:female) were similar across all areas, but the closed area had more females with eggs (8/10 females) than fished areas (1/17). Lamlash Bay marine reserve (2.67 km2) was established in 2008. In 2012–2015, crustaceans were sampled in summer inside the reserve, in three nearby (<2.5 km away) and four distant (10–20 km away; in 2013–2015 only) fished areas outside the reserve, using baited commercial shellfish pots deployed for 48–72 h (three rows of five pots/area) at 0–10 m depth. Organisms caught in pots were identified and counted. Abundance was derived from catch/unit effort. All lobsters, brown crabs, and velvet crabs were measured (carapace length), sexed, and fecundity recorded.Study and other actions tested