Action

Cease or prohibit all types of fishing in a marine protected area

How is the evidence assessed?
  • Effectiveness
    not assessed
  • Certainty
    not assessed
  • Harms
    not assessed

Study locations

Key messages

  • Seventy-nine studies examined the effects of ceasing or prohibiting all types of fishing in a marine protected area on fish populations. Fifteen studies were in the Indian Ocean (Kenya, Tanzania, South Africa, Mozambique, Madagascar, multiple African countries, Australia). Twelve studies were in the Mediterranean Sea (Spain, France, Italy). Ten studies were in the Pacific Ocean (New Zealand, USA, Hawaii, New Caledonia, Costa Rica, Tonga, Vanuatu, Solomon Islands). Seven studies were in each of the Coral Sea (Australia, Vanuatu), the Tasman Sea (New Zealand, Australia) and the Atlantic Ocean (Brazil, USA, Puerto Rico, Argentina, South Africa, UK, Canary Islands, Portugal, Turks and Caicos Islands). Four studies were in the Philippine Sea (Philippines). Three studies were in the Caribbean Sea (Belize, Puerto Rico). One study was in each of the Gulf of Mexico (USA), the Java Sea (Indonesia), the Pacific and Indian Oceans (multiple countries), the Sulu Sea (Malaysia) and the North Sea (Norway). Six studies were reviews of marine reserves (New Zealand, Latin America/Caribbean, regions unspecified and across the world).

 COMMUNITY RESPONSE (26 STUDIES) 

  • Community composition (7 studies): Seven site comparison studies (two replicated, and one before-and-after) in the Mediterranean Sea, Indian Ocean, Philippine Sea and the Atlantic Ocean found that protected areas where all fishing had been prohibited for between three and 16 years, had a different fish community composition, compared to fished areas. 
  • Richness/diversity (22 studies): Fourteen of 20 site comparison studies (eight replicated, one replicated and paired, and one before-and-after) in the Indian Ocean, Mediterranean Sea, Philippine Sea, Tasman Sea, Atlantic Ocean, Caribbean Sea, Coral Sea and the Pacific Ocean, found that marine protected areas that had prohibited all fishing for between one to more than 25 years, had higher fish species/richness compared to fished areas. Six studies found similar fish species/richness between one and 20 years after all fishing was banned in protected areas, compared to fished areas. One systematic review in the Atlantic and Pacific Oceans found no difference in species richness between unfished protected areas and fished areas. One replicated, site comparison study in the Indian Ocean found that the effects of prohibiting all fishing on fish species richness/diversity after 15 years varied with the sampling method used.

POPULATION RESPONSE (66 STUDIES)

  • Abundance (64 studies): Thirty of 54 site comparison studies (18 replicated, eight replicated and paired, two before-and-after, one paired and before-and-after, and one replicated and before-and-after) in the Indian Ocean, Atlantic Ocean, Mediterranean Sea, Pacific Ocean, Tasman Sea, Coral Sea, Philippine Sea, Caribbean Sea, Gulf of Mexico, and the Sulu Sea, found that marine protected areas that had been prohibiting all fishing for up to 25 years or more, had higher abundances (density and/or biomass) of all fish (total fish biomass, total fish density), fishery targeted fish species, non-fishery targeted fish species and all or most of the individual fish species/groups monitored, except fish densities (all or most) and non-fishery targeted species, compared to unprotected fished areas and/or partly-fished protected areas. The studies also found that in some cases where the total fish biomass or densities were higher in no-fished areas, the effect varied between individual groups of fish based on species family and/or position in the food chain, commercial target and non-target species, fish sizes, depth and habitat types. Eight studies found that inside protected areas prohibiting all fishing there were similar abundances of all fish, and all or most of the individual fish species/groups monitored, compared to fished areas between one and 20 years after implementation. The other sixteen studies found that the effect of prohibiting fishing in protected areas for three to 20 years on fish abundance varied between fish species or groups and on their fished status (fishery target or non-target) and/or position in the food chain. One also found that the effect varied with size or age of the protected areas. Five of six reviews (three systematic) across the world, in the Pacific and/or Atlantic Oceans and in unreported regions found that non-fished marine reserves with one to 27 years of protection had higher abundances of all fish, all fish and invertebrates combined and blue cod compared to fished areas, but there were differences between species/groups and fishing intensity outside reserves. The other review found that fish abundance varied between species in no-take marine reserves between one and 25 years old, and was affected by food chain position, level of exploitation and duration of protection. One replicated study in the Pacific Ocean found a long-term decline in the abundance/presence of eight of 12 shark and ray species inside an established (>15 years) no-fishing protected area, however enforcement was poor. One before-and after, site comparison study in the Pacific Ocean, found no differences in overall fish abundance between a marine reserve closed permanently to fishing for five years and a closed area that was harvested for two years during the same period. One site comparison study in the Coral Sea found that in a no-take zone of an area protected for at least 10 years, fish abundance of four of six fish groups were similar to no-entry and fished zones, but two had lower abundance than the no-entry zone. One replicated, paired, site comparison study in the Tasman Sea found that in a non-fished marine park zone abundance of commercially targeted fish was higher than partly fished zones but lower than unprotected areas after four to eight years.
  • Reproductive success (1 study): One site comparison study in the Mediterranean Sea found more eggs of four commercially targeted fish species inside a non-fished marine reserve enforced for three years than in fished areas outside the reserve.
  • Survival (1 study): One site comparison study in the Atlantic Ocean found that prohibiting all fishing in a marine protected area for three years resulted in similar survival of red hind grouper, compared to fished areas.
  • Condition (20 studies): Two global review studies (one systematic) and two systematic reviews in the Pacific Ocean and the Atlantic and Pacific Oceans found that prohibiting all fishing in marine protected areas for one to 27 years resulted in larger fish overall and larger blue cod compared to fished areas, but there were differences between individual fish families or species. Eight of 11 site comparison studies (four replicated, one before-and-after, one paired, and one replicated and paired) in the Tasman Sea, Pacific Ocean, Indian Ocean, Mediterranean Sea, Atlantic Ocean, Java Sea and the Philippine Sea, found that non-fished protected areas had larger fish overall and larger individuals of all or most of the fish species/groups monitored, compared to fished areas, after one to 22 years. The other three studies found similar fish sizes of all or all but one species, compared to fished areas one to 16 years after all fishing was prohibited. Three site comparison studies (one replicated) in the Coral Sea, Caribbean Sea and the Atlantic Ocean found that fish size in protected areas that had not been fished for six to more than 20 years, varied between fish species or food chain groups. One site comparison study in the Atlantic Ocean found that red hind grouper were larger, but had similar growth, in an area protected from fishing for three years compared to fished areas. One site comparison study in the Atlantic Ocean found that young lemon sharks in areas protected from fishing for 20 years had similar growth rates, but lower condition, than sharks in unprotected fished areas.

BEHAVIOUR (2 STUDIES) 

  • Behaviour change (2 studies): One replicated, site comparison study in the Pacific and Indian Oceans found that surgeonfish and parrotfish inside established protected areas where fishing was prohibited, showed a similar avoidance response to fishing gears as in fished areas, and this increased with increasing fishing intensity outside the protected areas. One replicated, site comparison study in the Indian Ocean found that in non-fished areas protected for one and 24 years, fish grazing rates were higher compared to fished areas.

 OTHER (15 STUDIES)

  • Catch abundance (2 studies): One of two site comparison studies in the Mediterranean Sea and Pacific Ocean found that commercial fish catch rates in small-scale traditional fisheries were highest closest to a marine reserve closed to all fishing for 22 years, and decreased with increasing distance from the reserve. The other study found that there was no increase in fish catch rates in commercially landed catch in the five years after a no-fishing zone was implemented in a co-managed protected area.
  • Stock biomass (1 study): One replicated, site comparison study in the Indian Ocean found that the stock biomass (the harvested portion of the population) of reef fish species was highest in enforced protected areas closed to all fishing, compared to various other area management regimes. 

 

About key messages

Key messages provide a descriptive index to studies we have found that test this intervention.

Studies are not directly comparable or of equal value. When making decisions based on this evidence, you should consider factors such as study size, study design, reported metrics and relevance of the study to your situation, rather than simply counting the number of studies that support a particular interpretation.

Supporting evidence from individual studies

  1. A review in 1993 of 11 case studies of reef marine reserves across the world prohibiting all types of fishing (Roberts & Polunin 1993) reported that most had increased abundance and size of fish between one and 15 years after protection compared to adjacent fished areas, but there were differences between species or family groups and level of exploitation, and with depth and fishing intensity outside the reserves. Three of four no-take reserves in the Philippines had higher overall fish abundances than fished areas after only one year and in one it had doubled after eight years closure (data not reported). For two reserves in the Caribbean Sea (Belize and Netherland Antilles) established for four years, higher fish densities and average sizes were found for a small number of species, but the biomass of four of five commercially targeted family groups was greater (unfished: 0.1–6.0 kg/count, fished: 0.1–2.0 kg/count), however the effect differed with depth and level of fishing intensity in adjacent fished areas. In a reserve in the Red Sea (Egypt), the effect of prohibiting fishing for 15 years on the abundance and biomass of commercial species was variable and for seven species where they were higher in reserves, average fish weights increased with decreasing fishing intensity (unfished: 42–1,333 g, lightly fished: 41–678 g, fished: 19–447 g). Field studies of reserves were reviewed (search methods not described) and results from sites throughout the tropics discussed.

    Study and other actions tested
  2. A site comparison study in 1992–1994 of two protected coral reef areas in the Indian Ocean, Kenya (Watson et al. 1996) found that prohibiting all fishing in a marine park for up to five years resulted in higher abundances of five of six fish family groups compared to a nearby marine reserve that permits traditional fishing types, but diversity was similar. The abundance of two of three non-commercially fished family groups were higher at the non-fished reef (butterflyfishes Chaetodontidae, non-fished: 52–58, fished: 23–40 fish/transect; damselfishes Pomacentridae, non-fished: 769, fished 412 fish/transect) and wrasses Labridae were similar (non-fished: 56, fished: 52 fish/transect). Abundances of commercially fished groups (emperors Lethrinidae, snappers Lutjanidae and groupers Serranidae) were greater at the no-fishing reef than the fished reef (data were not statistically tested). No differences in fish species number and diversity between non-fished and fished reefs were found (data reported as statistical results). In September-October 1992 and January-March 1994, visual underwater censuses (250 × 10 m transects) of six representative reef fish families were done at a series of sites (number was not reported) at both the Kisite Marine National Park (closed to all fishing types) and Mpunguti Marine National Reserve (traditional fishing such as hand lining and basket trapping only is permitted). Enforcement began in both areas in 1989.

    Study and other actions tested
  3. A replicated, site comparison study in 1997 of two areas of coralline flats in the Pacific Ocean, northern New Zealand (Cole & Keuskamp 1998) found that protected areas where all fishing is prohibited had a higher overall density of fish that feed on urchins Evechinus chloroticus, and they were larger, compared to adjacent fished areas after 22 years. Data were not tested statistically. The total number of urchin-eating fish was greater in no-fishing areas (161) than fished (76) and they were of larger sizes (data presented as length frequencies). Individually, the densities of four of the eight species were higher in non-fished areas, one was the same and three had very low densities in both areas (see paper for individual data). In December 1997, eight potential fish predators of sea urchins were surveyed by underwater visual census (10 replicates of 25 × 5 m transects) at five sites in Cape Rodney-Okakari Point Marine Reserve (no-take since 1975) and five fished sites in an adjacent area.

    Study and other actions tested
  4. A replicated, site comparison study in 1997 of two areas of sand and reef in the Tasman Sea off northeast New Zealand (Babcock et al. 1999) found that prohibiting all fishing in two adjacent marine reserves established for 15 and 22 years, resulted in an increased abundance and size of snapper Pagrus auratus compared to adjacent fished areas outside the reserves. Across both reserves, abundances of snapper above the minimum legal length (270 mm) were higher in no-take reserves than fished areas (no-take: 2–5 fish/30 min, fished: <1 fish/30 min) and average total length was greater (no-take: 327 mm, fished: 191 mm). Snapper, as the most abundant predatory fish, were surveyed in October and November 1997 at two no-take reserves: the Leigh Marine Reserve (549 ha, established 1975) and Tawharanui Marine Park, 15 km to the south (350 ha, established 1982). Maximum numbers and estimated length of snapper responding to remotely deployed baited video camera deployments of 30 minutes were recorded. Four replicate deployments were made at six sites inside and six outside the Leigh reserve, and three both inside and outside Tawharanui (72 deployments of 30 mins at 16–24 m depth).

    Study and other actions tested
  5. A replicated, site comparison study in 1996 of fifteen patch coral reef sites in the Indian Ocean off east Africa (McClanahan et al. 1999) found that prohibiting all fishing in two marine parks for 5 and 22 years resulted in higher total fish biomass and similar fish densities and total species richness compared to unprotected fished reef areas, and there were differences between individual fish family groups. Across both parks, total fish biomass was higher at non-fished (806 kg/ha) than fished (230kg/ha) reefs and individually for seven of eleven fish groups (see paper for data by group). There were no differences in average densities of eight of eight fish groups between non-fished and fished reefs (non-fished: 0–317 fish/500 m2, fished: 0–609 fish/500 m2), although higher densities were recorded in non-fished reef areas for 26 of 134 individual species and a lower density for one. Overall species richness was similar at non-fished (40–50 fish/500m2) and fished (28–39 fish/500m2) reefs, and higher at non-fished reefs for four of eight family groups (see paper for data by group). Fish were surveyed at five non-fished and ten fished coral reefs sites off southern Kenya and Tanzania (sampling times were not reported). Three non-fished reefs were in the Kisite Marine National Park (10 km2, no-take since 1974) and two in the Chumbe Island Coral Park (500 m stretch of reef, no-take since 1991). At each reef site the fish assemblage was quantified along two 5 x 100 m transects by two methods: one to estimate wet weight by family group and one to record the number of individuals per species and the number of species per transect. The authors used a non-standard threshold for statistical significance (0.07).

    Study and other actions tested
  6. A site comparison study in 1997–1998 of an area of rock and cobble in the Tasman Sea, South Pacific Ocean, off New Zealand (Cole et al. 2000) found that prohibiting all fishing activity for 4 years in a marine reserve did not result in higher overall abundances of blue cod Parapercis colias compared to adjacent fished areas outside, but blue cod inside the reserve were larger. Numbers of blue cod were similar inside the reserve (44 fish/transect) to fished areas (41 fish/transect) at all depths except 20 m. However, the lengths of blue cod inside the reserve were on average 4 cm longer (20–40 cm) than those found in commercially fished areas (21–25 cm). Blue cod were surveyed five times between January 1998 and 1999, at two sites inside the Long Island–Kokomohua Marine Reserve, Marlborough Sounds (619 ha, established as no-take in 1993) and two adjacent (2–4 km apart) fished sites where commercial fishing for blue cod is prohibited but recreational fishing effort can be high. At each site, numbers of cod were recorded during four minute–long diver visual censuses at depths of 5, 8, 11, 14, 17 and 20 m. At each depth, lengths of blue cod were estimated in two-minute time intervals, along a 2 m wide transect.

    Study and other actions tested
  7. A systematic review in 2000 of 24 studies of marine reserves across the world (Mosquera et al. 2000) found that overall fish abundance was higher, and fish were larger, inside no-take (all fishing types prohibited) reserves with 1 to 26 years of protection, compared to fished areas outside reserves. Overall fish numbers were on average 3.7 times higher inside non-fished reserves than outside. Abundances of fishery targeted species were higher in non-fished reserves than fished areas outside, but non-target species abundance was similar (data reported as statistical model results). Across all species, the effect of protection status (non-fished versus fished) on abundance depended on fish body size; the largest species were over 300% more abundant inside reserves and the effect increased with body size (data reported as model results). The systematic review used data from 24 studies to assess the effects of banning fishing in marine reserves. Twelve of the studies met the criteria for quantitative meta-analysis, the other 12 studies were not included in the meta-analysis.

    Study and other actions tested
  8. A replicated, site comparison study in 1998 of eight rocky and sandy sites in the San Juan Archipelago, northwest Pacific Ocean, USA (Tuya et al. 2000) found no differences in the abundances of copper rockfish Sebastes caurinus, quillback rockfish Sebastes maliger, China rockfish Sebastes nebulosus and lingcod Ophiodon elongatus between three marine research areas established for eight years where all fishing was prohibited, two areas protected for one year where only fishing for salmon was permitted, and sites open to fishing. Fish abundance data were not provided (reported as statistical results). The authors suggest the lack of increase in fish abundances inside protected areas was likely due to a lack of compliance and enforcement of the restrictions. In July 1998, three marine research reserves (established in 1990 and prohibiting all extractive activities except controlled research collection; sea urchin fishery closed since the late 1970s), two marine protected areas (designated in 1997; voluntary no-take zones where no finfish except salmon can be taken), and three unprotected areas were surveyed. Two divers identified and counted fish along 300 m2 transects on reef slopes up to 20 m deep (4 transects/site).

    Study and other actions tested
  9. A before-and-after, site comparison study in 1992–2001 of an area of shallow rocky reef in the Tasman Sea, New Zealand (Davidson 2001) found that the average density, size and catch abundance of blue cod Parapercis colias increased inside a marine reserve in the eight years after all fishing was prohibited, compared to before and recreationally fished areas outside. Cod density was higher inside the unfished reserve than outside from two years after closure (1995) compared to before (1995, inside: 3.2, outside: 1.9 fish/60 m2; 2001, inside: 6.5, outside: 2.9 fish/60 m2). Across all years following closure (1993–2000), average length of blue cod was higher in the unfished reserve than fished areas (data reported as statistical model results), and increased over time inside the reserve (2000: 265 mm, 1993: 228 mm) while outside lengths decreased (2000: 71 mm, 1993: 154 mm). Over the same period, experimental catch rates were higher and increased over time inside the reserve compared to outside (data reported as statistical model results). Long Island-Kokomohua Marine Reserve (619 ha) in the Cook Strait was designated as no-take (no-fishing) in April 1993. Blue cod numbers were surveyed annually from March 1992 to April 2001 by underwater visual transects (2 × 2 × 30 m), inside (four/five sites) and outside (four sites) the reserve. Size and catch rates were monitored from September 1993 to April 2000 at three sites inside and six outside the reserve using experimental baited hook and line fishing.

    Study and other actions tested
  10. A site comparison study in 1994–1997 of a surf-zone area in the Indian Ocean off the coast of South Africa (Cowley et al. 2002) found that four of four important shore-angling fish species were larger and more abundant in a marine park where all fishing was prohibited for over 16 years, compared to openly fished areas. Average fork lengths were greater inside the no-fishing marine park than fished areas for blacktail bream Diplodus sargus capensis (unfished: 284 mm, fished: 226 mm) zebra bream Diplodus cervinus hottentotus (unfished: 303 mm, fished: 248 mm) and galjoen Dichistius capensis (unfished: 365 mm, fished: 327 mm), and were similar for bronze bream Pachymetopon grande (unfished: 358 mm, fished: 354 mm). In addition, catch rates for all species were higher in the marine park (unfished: 4–13, fished: <1–3 kg/100 angler hrs; data were not statistically tested). Fish data were collected from monthly research shore-angling between February 1995 and January 1997 in the Tsitsikamma National Park (80 km of coastline where all fishing is prohibited; shore-angling banned since 1978) and in fished areas extending either side of the park from roving surveys of recreational shore- angler catches between April 1994 and February 1996.

    Study and other actions tested
  11. A replicated, paired, site comparison study in 2001–2002 in two coral reef areas of the Great Barrier Reef Marine Park, Coral Sea, Australia (Graham et al. 2003) found that prohibiting all fishing in no-take zones resulted, after 14 years, in a decline in abundance of six of nine fish species that are prey for the fishery targeted coral trout Plectropomus spp. compared to fished zones, while the biomass of coral trout was higher. Average prey fish densities were lower in the no-take than fished zones for six of nine species (no-take: 8–342 fish/site, fished: 12–507 fish/site) and were similar for three (see paper for individual species data). In addition, overall coral trout biomass was greater in the no-take zones (9,790 g/1,500 m2) than the fished zones (3,420 g/1,500 m2). Fish data were collected in two areas of the Great Barrier Reef Marine Park using five, 50 × 6 m, belt transects at each site: the Whitsunday Island group was surveyed in December 2001 at eight sites in no-take zones (no fishing permitted, 14 years of protection) and eight in fished zones; and the Palm Island group was surveyed in April 2002 at eight sites in no-take zones (14.5 years of protection) and eight sites in fished zones. Sites were at least 100 m apart.

    Study and other actions tested
  12. A replicated, paired, site comparison study in 2001–2002 of three coral reef areas in the Great Barrier Reef Marine Park, Coral Sea, Australia (Evans & Russ 2004) found that prohibiting all fishing in protected areas resulted in a higher biomass and density of two fish species targeted by commercial line fisheries after 14 years, and similar densities of two fish species not targeted by commercial fisheries, compared to nearby fished areas. Biomass and density in the no fishing areas was higher for the commercially fished coral trout Plectropomus spp. >35 cm (biomass: 6.6 kg/1,000 m2; density: 3 fish/1,000 m2) and Spanish flag snapper Lutjanus carponotatus >25 cm (biomass: 5 kg/1,000 m2; density: 14 fish/1,000 m2) than fished areas (coral trout: biomass; 1.3 kg/1,000 m2, density; 1 fish/1,000 m2; Spanish flag snapper: biomass; 2 kg/1,000 m2, density; 1 fish/1,000 m2). The density in the no fishing areas was not significantly different for the non–fished species scribbled rabbitfish Siganus doliatus (9 fish/1,000 m2) and golden butterflyfish Chaetodon aureofasciatus (19 fish/1,000 m2) compared to fished areas (rabbitfish; 13 fish/1,000 m2, butterflyfish: 19 fish/1,000 m2). Fish counts and size estimates were recorded by underwater visual surveys between December 2001–October 2002 at three no fishing reserves around the Palm, Whitsunday and Keppel Islands (spanning 600 km of the Great Barrier reef, no fishing for 14 years). Five replicate 50 × 6 m transects were randomly selected at six to 12 sites per protected and fished area, 200–400 km apart from each other.

    Study and other actions tested
  13. A review in 2004 of 20 studies of marine reserves across the world (Micheli et al. 2004) found that fish abundance and biomass in no-take marine reserves where all fishing was prohibited for between one to 25 years, varied between species compared to fished reference sites outside reserves, and the response was influenced by food chain position, level of fishing exploitation and duration of protection. Between 5 and 91% of fish species showed strong increases in abundance in no-take reserves compared to fished reference conditions, and 0–36% decreased in abundance. Where there were differences, greater abundances in no-take reserves were found to be associated with five of six food chain groups and for species targeted by fishing or the aquarium trade, with no overall response for non-targeted fish (data reported as response ratios and statistical results). Variation in species responses was also found with time since protection, with abundances of top predators increasing gradually and accounting for greater proportions of the total biomass in the reserves (data reported as response ratios). A literature search for field studies examining the effect of prohibiting all fishing types in no-take reserves on fish communities was carried out. A meta-analysis of data from 20 studies conducted at 31 different locations in which fish abundance and/or biomass for more than 10 individual species had been compared to fished reference sites was done. All studies used visual census (belt transects and point counts) apart from one study that used trammel nets to collect the fish data.

    Study and other actions tested
  14. A site comparison study in 1994–1998 in an area of coral reef in the North Pacific Ocean, Hawaii (Wetherbee et al. 2004) found that the short and long-term movement patterns of tagged and tracked giant trevally Caranx ignobilis indicated that a marine reserve where all fishing was prohibited for over 30 years was used by only certain sizes of trevally, and there were frequent movements outside the reserve into fished areas where some were caught by fishers, thus it provided limited protection from fishing. Average size of trevally caught inside the reserve was 28 cm total length (range 14–43 cm) and 22 cm (range 16–37 cm) for those caught outside. Of 289 conventionally tagged trevally 33 fish (11%) were recaptured after an average time at liberty of 346 days (min 2 d, max >7 y). A high percentage (79%) of the recaptured trevally were originally tagged inside the reserve, but only 15% were both tagged and recaptured there, while nearly one third were caught by fishers over 3 km away (up to 70 km). The movement activity of five fish tracked for 9–125 hours showed they spent considerable time inside the reserve but also made frequent movements outside (data reported as minimum convex polygons and kernel home range). Coconut Island (situated on 137,000 m2 of reef flat, 2.4 km linear perimeter) has been a marine reserve for over 30 years, with a no-fishing zone extending 8 metres seaward from the reef edge. Giant trevally sizes in and around the reserve were collected opportunistically throughout the year between 1994 and 1998 from research fishing. Long-term movements were monitored by recaptures over 9.5 years (dates of tagging were not reported) of 58 conventionally tagged trevally caught by rod and line inside the reserve and 231 caught by traps outside. The short-term movements of five trevally fitted with transmitters were tracked by boat using a hydrophone for periods up to 14 days (sampling times were not reported).

    Study and other actions tested
  15. A replicated, paired, before-and-after, site comparison study in 1983–2000 at two coral reef areas in the Coral Sea, Australia (Williamson et al. 2004) found that prohibiting all fishing (no-take) in two marine reserves resulted in an increase in density and biomass of coral trout Plectropomus spp. in the period from 3–4 years before establishment to 12–13 years after and compared to fished areas, and higher densities and abundances of fishery targeted species, but not non-target species compared to fished areas 12–13 years after. At both no-take reserves, the average density and biomass of targeted coral trout Plectropomus spp. was higher (1999–2000, density: 7–17 fish/1,000 m2; biomass: 12–16 kg/1,000 m2) than in pre-protection (1983–1984, density: 2–3 fish/1,000 m2; biomass: 2 kg/1,000 m2) and fished areas (1999–2000, density: 3–5 fish/1,000 m2; biomass: 3 kg/1,000 m2), the latter two areas being similar. In 1999–2000, average coral density and biomass of a second targeted fish, stripy sea perch Lutjanus carponotatus was higher in both reserves than fished areas (density: 12–23 vs 7 fish/1,000 m2, biomass: 4–5 vs 2 kg/1,000 m2) but average density and biomass of non-target fish did not differ (density: 56–86 fish/1,000 m2, biomass: 7–17 kg/1,000 m2). Reef fish were surveyed by underwater visual census at two island group marine reserves in the Great Barrier Reef Marine Park (no fishing since 1987). In the period before protection, five replicate transects (50 × 20 m) were done in 1983 (one reserve only, 2 sites) and 1984 (both reserves, 2 sites each). In November 1999 to June 2000, no-take and fished zones at both reserves were surveyed by five replicate 50 × 6 m transects (9–12 transects/no-take and fished areas).

    Study and other actions tested
  16. A site comparison study in 2002–2003 of three areas of artificial rock in the Adriatic Sea, Italy (Guidetta, Bussotti & Boero 2005, same experimental set-up as Guidetta et al. 2005) found higher abundances of white seabream Diplodus sargus, two-banded seabream Diplodus vulgaris and gilt-head seabream Sparus aurata at a breakwater in a marine protected area where all fishing had been prohibited for 16 years, and there were more medium and large individuals, compared to two nearby fished breakwaters. The density of white (unfished: 5.0–7.8, fished: 0.0–1.7 fish/125m2) and two-banded seabream (unfished: 11.6–45.7, fished: 1.0–14.3 fish/125 m2) was higher at the unfished breakwater than fished ones in two of the four sampling times, and all but small individuals were more abundant (white, small: 0.5 vs 0.0–0.5, medium: 3.7 vs 1.7–1.8, large: 1.8 vs 0.1–0.3; two-banded, small: 0.0 vs 0.0–1.0, medium: 11.1 vs 3.4–3.8, large: 2.2 vs 0.2–0.3 fish/125m2). Gilt-head seabream were present only at the unfished breakwater in three of four sampling times and were more abundant in the other (unfished: 1.3–2.2, fished: 0.5 fish/125m2). Fish were surveyed at one breakwater in the Miramare Marine Protected Area (121 ha, no fishing since 1986) and two fished breakwaters (adjacent and 3 km away) four times between spring 2002 to summer 2003. Four underwater visual transects (25 × 5 m) were done at each breakwater. The breakwaters were transplanted boulders 1–3 m wide running parallel to the coast, extending from the surface to depths of 5–8 m.

    Study and other actions tested
  17. A site comparison study in 2002–2003 of three areas of artificial rock in the Adriatic Sea, Italy (Guidetta et al. 2005, same experimental set-up as Guidetta, Bussotti & Boero 2005) found that a breakwater in a marine protected area where all fishing was prohibited for 16 years had a different fish assemblage, a higher species richness, and a similar total fish density but higher density of commercially targeted fish species, compared to two unprotected fished breakwaters. The fish assemblage at the unfished breakwater differed to both fished breakwaters in three of four sampling times, and only one in the final sampling time (data reported as statistical results and graphical analysis). In all four sampling times, species richness was higher at the unfished breakwater (24–27) than fished ones (13–22). Overall fish density was higher at the unfished breakwater in only one of four sampling times, however the individual densities of eight of 12 commercially targeted species were higher at the unfished breakwater in two or more sampling times, and schooling fish density was higher in all four sampling times (data reported as statistical results). The Miramare marine protected area was designated in 1986 and a fishing ban is successfully enforced. Four surveys using two different methods were undertaken from spring 2002 to summer 2003 at one breakwater in the Miramare Marine Protected Area (121 ha, no fishing since 1986) and two fished breakwaters (adjacent and 3 km away). Each sampling time, four transects (25 × 5 m) and four, point counts (5 m radius) were done per breakwater. The authors noted differences in the data between the two census methods.

    Study and other actions tested
  18. A replicated, paired, site comparison study in 2002 of four coral reefs off two islands in the Bohol Sea, Philippines (Abesamis et al. 2006) found that prohibiting all types of fishing resulted in greater abundance and biomass of commercially targeted fish at one of two marine reserves established for 15–20 years, and higher abundance of non-target fish, compared to nearby fished areas. Abundance and biomass of commercially targeted fish were higher inside Apo marine reserve compared to fished areas (density, inside: 68, outside: 26 fish/500 m2; biomass, inside: 90, outside: 25 kg/ 500 m2) and were similar inside and outside Balicasag reserve (inside: 44, outside: 34 fish/500 m2; biomass data not reported). The abundance of non-commercially targeted fish was greater inside both marine reserves than fished areas (inside: 75–129 fish/ 250 m2, outside: 90–147 fish/250 m2). In November and December 2002, fish were surveyed at one site inside and one outside each of the Apo (450 m length of reef, no-take since 1982) and Balicasag marine reserves (650 m long reef, no fishing since 1985, the collection of deep–water ornamental shells is permitted). Fish were surveyed along fifteen 50 × 10 m transects/site: commercial fish 5 m either side (96 species from 13 families) and non–commercial fish 2.5 m either side of the transects (four species of damselfish Pomacentridae, 15 species of butterflyfish Chaetodontidae).

    Study and other actions tested
  19. A site comparison study in 2001–2003 of a reef archipelago in the Atlantic Ocean, Brazil (Garla et al. 2006) found that young Caribbean reef sharks Carcharhinus perezi were more abundant inside a marine protected area where fishing had been prohibited for over 12 years, compared to an adjacent fished area. Average catches of sharks were higher inside the unfished area than the fished (unfished: 0.16 sharks/h, fished: 0.03 sharks/h). Sharks caught in both areas were almost all smaller immature individuals (71–170 cm). Fernando de Noronha Archipelago (26 km2) is 345 km off the northeastern coast of Brazil and has a marine protected area, no fishing since 1988, around the coastline of its main island out to 50 m water depth. The rest of the area allows fishing and boat traffic. Monthly from March 2001 to February 2003, fishing for sharks was done at 148 randomly selected sites around the archipelago, inside (79) and in the fished area outside (69) the protected area. At each site two baited, single-hook handlines were deployed simultaneously from a small boat. Catch per unit effort of sharks was calculated from the time the first hook was deployed to the time the last hook was removed. Number, length, and sex of captured sharks (143) were recorded.

    Study and other actions tested
  20. A replicated, site comparison study in 1996–2004 at seven coral reef sites in the Indian Ocean off Kenya and Tanzania (McClanahan et al. 2006) found that in a large permanent no-take zone of a marine protected area where fishing was prohibited for over 20 years, there was higher total fish biomass and species richness, but biomass varied between fish family groups, compared to reefs managed collaboratively for less than 10 years by gear restrictions and temporary fishing closures. Total weight of fish was greater in the area with a permanent no-take zone compared to without in two of two years sampled (weight: 682–1,354 vs 260–457 kg/ha) but the responses differed by individual fish family group (see paper for data), and total number of fish species was higher (with: 47–51, without: 38–41 species/500 m3). Data were collected from sites in two locations: three reefs in a 10 km2 area of the Kisite-Mpunguti Marine National Park in Kenya (established 1973) permanently closed to all extractive activities and adjacent to a gear-managed reserve; and four small reefs (0.25–3.0 km2) in the Mtang’ata Collaborative Management Area in Tanzania (established in 1996) managed by gear restrictions and small voluntarily and temporary closed areas (some illegal fishing reported). At each reef site, fish communities were surveyed twice (in 1996 and 2003–2004) by underwater visual census along two 5 × 100 m belt transects at each site.

    Study and other actions tested
  21. A site comparison study in 2004–2005 of a rocky reef island in the Tyrrhenian Sea, off Italy (Cecchi et al. 2007) found that prohibiting all fishing in a marine protected area for nine years resulted in a different overall fish assemblage compared to a recreationally fished area, and the abundance of recreationally targeted species was higher at the deeper of two depths. The overall fish assemblage was different between the unfished and fished areas at 5 m and 20 m depths (reported as statistical results). Average number of individuals of species targeted recreationally was higher in the unfished area (9) than in the fished (6) at 20 m depth, but similar at 5 m depth (unfished: 4, fished: 3). Fish were surveyed along 11 km of coastline around Giannutri Island in areas with two different protection levels (established approximately 1993): one where all human activity is banned, and one where commercial fishing is banned but recreational fishing and other activities are permitted. In July and September 2004 and March and May 2005, fish were sampled by visual census at two sites/protection level at 5 m and 20 m depth. Fish within an imaginary cylinder 5 m high and 10 m in diameter were recorded.

    Study and other actions tested
  22. A replicated, site comparison study in 2004 of five coral reef sites in the Florida Keys, Atlantic Ocean, USA (Kramer & Heck 2007) found that prohibiting all fishing within marine protected areas (no-take) for 6 years resulted in higher biomass, body length and abundance of some reef fish species and sizes, but not others, compared to unprotected fished reefs. The average biomass of one of two species of groupers Serranidae spp. and one of three snappers Lutjanidae spp. was higher inside (grouper: 1,190; snapper: 910 g/125 m2) than outside no-take areas (grouper: 130; snapper: 30 g/125 m2), but was similar for the others (inside: 590–2,400, outside: 100–2,500 g/125 m2; see paper for individual data). Average body lengths of two of the three snappers were greater in no-take areas, while no differences were found for the other snapper and the only grouper for which there was sufficient data (data reported as statistical results). For three groups of herbivorous fish (see original paper for species), adult sizes of two were more abundant in no-take areas (inside: 0.30–0.98, outside: 0.13–0.74 m2) and abundances of immature sizes were lower (inside: 0.04–0.60; outside: 0.12–1.50 fish/m2), while abundance of the other species was similar for both adults and immature fish (inside: 0.05–0.30, outside: 0.03–0.10 fish/m2). Patch reefs were sampled in three Special Protected Areas (average 0.5 km2, established 1997, no resource extraction) and at two fished reefs (1 to 3 km apart). Predatory and herbivorous fish were recorded along three 25 × 5 m and 20 × 1 m belt transects, respectively. Predatory fish were surveyed on 5-6 days in June-September 2004 and herbivorous fish on 7–9 days in June-September 2003 and 2004.

    Study and other actions tested
  23. A replicated, site comparison study in 1998–2004 of five coral reefs in the Philippine Sea, Philippines (Samoilys et al. 2007) found that, over six years, no-take marine reserves in which all fishing had been prohibited for at least one to three years, had different fish communities compared to adjacent and distant fished areas outside, fish abundances varied between species and level in the food chain, and the differences were greater at reserves with the highest enforcement and compliance history. Fish communities differed between all areas (non-fished, adjacent fished and distant fished) and differences between non-fished and adjacent fished areas were strongest at the two of five reserves with the strictest protection levels (data reported graphically and as statistical results). For fish species at the top of the food chain, abundance was higher at two of the five non-fished reserves than adjacent and distant fished areas across all years (non-fished: 4–28, fished: 3–34 ind/250 m2), and varied between areas at the other reserves over time. Density of fish species in the middle of the food chain was similar between sites (non-fished: 0–148, fished: 0–151 ind/250 m2). For the dominant fish group at the bottom of the food chain Pomatocentridae, density was higher in non-fished areas than fished for two reserves, one with good enforcement (non-fished: 7–149, fished: 0–70 ind/250 m2), and density did not differ between areas at the other three reserves. In addition, the response to no fishing varied between individual fish families and abundances of larger and/or targeted fish by fishers was generally higher inside the reserves, while non-preferred species were more abundant outside. Data was collected at five no-fishing reserves in the Bohol Strait, differing in size (11–50 ha), age (established 1995–1999) and history of enforcement and compliance. One site inside and one outside (within 1,000 m) each of the reserves and at three distant fished sites were monitored twice a year in February-May and August-November from 1998-2004. Fish were surveyed by underwater visual censuses along four 50 x 5 m transects at each site. Fish were counted, fish length measured, and identified to species family and food chain group.

    Study and other actions tested
  24. A replicated, site comparison study in 2001–2005 of four coral reefs on Abrolhos Bank, South Atlantic Ocean, Brazil (Francini-Filho & De Moura 2008) found that prohibiting all fishing in two no-take reserves, protected from 0 and 18 years, resulted in higher biomass of commercially targeted and non–targeted fish at the older reserve compared to multiple use protected areas and unprotected openly fished areas, but the response varied with fish species and/or level in the food chain. Across all years, total biomass of both commercially targeted and non-target fish groups was higher in the older no-take reserve than any other area, but openly fished areas had higher biomass than the younger and multiple use reserves (data reported as statistical results). However, the response varied by fish species and/or food chain group (see paper for individual data). Reef fish were monitored annually in January-March 2001–2005 in four areas: one reserve area in the Abrolhos National Marine Park (no-take since 1983); one no-take reserve (since November 2001) and one multiple-use area (co-managed since 2000, use permitted by locals only, zoning and gear restrictions) in the Corumbau Marine Extractive Reserve, and an unprotected fished, open access area. Some illegal poaching was reported in the no–take areas. Two habitats at three to seven sites were sampled in each management area by underwater visual census (15–20 samples/habitat/site/year). Fish were identified and counted in a 2 m or 4 m radius.

    Study and other actions tested
  25. A replicated, site comparison study in 1992–2004 of 12 coral reefs in the Indian Ocean, off Kenya (McClanahan 2008) found that grazing rates of fish on seagrass Thalassia hemprichii over a 12-year period were higher in no-take marine protected areas established for one and 24 years and where all fishing is prohibited, compared to fished reefs. For two different measures of fish grazing, both the rate of fish bites on seagrass (unfished: 53%, fished: 1%) and the average amount of algae eaten by fish at the unfished reefs (unfished: 65 kg/ha/d, fished: 2 kg/ha/d) were higher than fished reefs. In addition, coral cover in the unfished reefs (29%) was higher compared to the fished areas (16%). Fish grazing was monitored annually by two methods, at five sites in three marine protected areas: Malindi and Watamu (all fishing prohibited since 1968) and Mombasa (no-take since 1991); and seven sites in heavily fished areas. Firstly, thirty, 10-cm long blades of seagrass were soaked for 24 hours at each site and the numbers bitten by finfish recorded. Secondly, the biomass of selected fish herbivores along three to five, 500 m2 belt transects at each site was used to estimate the amount of algal biomass eaten per day (based on 16% of body weight) per wet weight of fish.

    Study and other actions tested
  26. A systematic review of unpublished data from 11 studies of five marine reserves surveyed between 1992–2002 in the southwestern Pacific Ocean around New Zealand (Pande et al. 2008) found that overall, blue cod Parapercis colias were found to be larger and more abundant in reserves where all fishing had been prohibited between 1 to 27 years, compared to fished areas outside. In nine of 10 cases, blue cod total length was greater inside unfished reserves (25–31 cm) than outside (20–27 cm) and for eight of 11 cases, cod were more abundant inside (0.003–0.099 fish/m2) than outside (0.003–0.051 fish/m2) unfished reserves. In addition, although the magnitude of the differences varied between reserves, blue cod length and abundance was not affected by the size or age of the reserves. A meta-analysis of 11 unpublished blue cod datasets from surveys of five no-take marine reserves was done. The reserves ranged in size from 93–2,400 ha and in age since protection from 1–27 years and prohibited all types of fishing. Blue cod length and abundance inside each reserve was compared to adjacent areas outside (distances apart were not reported).

    Study and other actions tested
  27. A site comparison study in 1992–2005 of three rocky areas in the northwest Mediterranean off the coast of Spain (Rius & Zabala 2008) found that banning all types of fishing for at least nine years in a marine protected area resulted in higher biomass of white seabream Diplodus sargus and gilthead bream Sparus aurata, compared to nearby recreationally fished only and unprotected fished areas. Across all years, the average biomasses of white and gilthead bream were higher in the unfished area (white: 19.1 g/m2, gilthead: 0.8 g/m2) than the other areas, and were similar between partially fished (white: 5.9 g/m2, gilthead: 0.1 g/m2) and fished areas (white: 6.1 g/m2, gilthead: 0.2 g/m2). Fish were sampled annually from 1992–2005 at three nearby sites, up to 2 km apart: a no-take reserve in the Medes Islands Marine Protected area (no extractive activities, since 1983); a partial reserve (angling permitted but no collection of subtidal animals since 1990); and a fished stretch of coastline. Numbers of fish at each site were recorded by underwater visual transects (no further sampling details were reported).

    Study and other actions tested
  28. A site comparison study in 2005–2007 of one coastal site in the Mediterranean Sea, France (Seytre & Francour 2008, same experimental set-up as Seytre & Francour 2009) found that prohibiting all fishing activity in a marine protected reserve increased total fish biomass, abundance and species richness compared to outside the reserve where fishing is allowed, one to four years after protection. Average total biomass, abundance, and species richness was higher at one of two sampling sites inside the reserve in 2006 and at both in 2007 (biomass, 2006: 3 kg, 2007: 6–10 kg; abundance, 2006: 6, 2007: 9–10; species richness, 2006: 3.5, 2007: 7) compared to sites outside the reserve (biomass, 2006: <1 kg, 2007: <1–2 kg; abundance, 2006: 1, 2007: 1–4; species richness, 2006: 1, 2007: 1–4). In addition, the commercial fish assemblage was different inside and outside the reserve in 2006 and 2007 but not in 2005, and no differences were found for assemblages of small-sized fish over seagrass Posidonia oceanica seabed. Fish were monitored inside and at two locations outside (adjacent areas north and south) the Cape Roux Marine Protected Area (450 ha, all fishing types prohibited since December 2003). In October 2006 and June 2007, six trammel net deployments sampled all fish at two sites inside the reserve and one in each location outside. From 2005–2007, at total of 28 commercial (sampled every season for 2.5 years) and 28 small fish species (sampled in spring for one year) were surveyed by underwater visual census six to 10 times at two sites in each location.

    Study and other actions tested
  29. A site comparison study in 2005–2007 of one no-take coral reef marine reserve, closed to fishing for 15 years, and two unprotected (fished) reefs in the Pacific Ocean off New Caledonia, France (Chateau & Wantiez 2009) reported that the tracked movements of four of four commercially important reef fish species indicated that most did not move from the unfished reef to the fished reefs and thus were largely protected from fishing, however, some fish did make large-scale movements outside of the reserve reef. Data were not tested for statistical significance. Of 45 fish tracked, a total of 36 (80%) did not move between the three reef sites but nine fish (20%), including at least one individual from each of the four species, moved distances of 510–6,000 m outside the reserve reef. The South Lagoon Marine Park was created in 1990 and has one temporary and nine permanent no-take marine reserves where all fishing is prohibited. From July 2005 to January 2007, movement data for 45 fish of four major commercial reef species (19 individuals of two Serranidae spp. and 26 individuals of two Scaridae spp.) fitted with transmitters were collected by 23 hydrophones deployed at 4–13 m depth around one reserve and two fished reefs. After being fitted with transmitters, 35 fish were released at their original site of capture (28 in the reserve and 7 in a fished reef) and ten fish that were caught in a fished reef were released inside the reserve. The number of days each fish was detected for was reported only for some individuals.

    Study and other actions tested
  30. A site comparison study in 2000–2004 of an area of reef in the Indian Ocean, off South Africa (Götz et al. 2009) found that prohibiting all types of fishing in a marine protected area for over 10 years resulted in a different fish community, higher diversity and a higher abundance of four of the eight most common fish species compared to an adjacent fished reef. Fish community composition was found to differ between unfished and fished areas using both sampling methods and diversity was higher in the unfished area for the visual census survey only (unfished: 1.7, fished: 1.5). For both sampling methods, frequencies of roman Chrysoblephus laticeps, steentjie Spondyliosoma emarginatum, dreamfish Sarpa salpa and blacktail Diplodus sargus capensis were higher in the unfished area (unfished: 8–55%, fished: 5–36%) and fransmadam Boopsoidea inornata, blue hottentot Pachymetopon aeneum, santer Cheimerius nufar and dageraad Chrysoblephus cristiceps were lower (unfished: <1–14%, fished: <1–42%; see paper for species individual data by method). Fish were sampled inside and outside the Goukamma Marine Protected Area (40 km2, all fishing prohibited since 1990) using two methods: standardised angling (111 sites inside, 162 outside) and underwater visual census (15 inside, 29 outside 44 sites). Angling surveys were carried out in all seasons from 2000 to 2003 by a team of 3–5 anglers. Seasonally in 2001–2004, fish were counted by divers in an area of up to 5 m radius.

    Study and other actions tested
  31. A site comparison study in 2002–2003 at six reef sites in the southwestern Atlantic Ocean, off Puerto Rico (Lopez-Rivera & Sabat 2009) found that prohibiting all fishing in a marine protected area for three years resulted in similar abundances of red hind grouper Epinephelus guttatus and its associated prey fish species, and larger red hind size but similar growth and survival, compared to adjacent fished areas. There were no differences in red hind density (unfished: 9–23, fished: 6–26 fish/ha), growth rate or average annual survival rate between management types, but average total length was larger inside the unfished area (data reported as statistical results). Average abundances of three fish species and six family groups that are prey for red hind were similar between areas (unfished: 0–44, fished: 0–46 fish/no. census; see paper for species individual data). Red hinds and the prey fish community were surveyed at three sites inside the Luis Peña Channel Marine Fishery Reserve (4.75 km2, established as no-take in September 1999) and three sites in adjacent fished areas outside. Fish abundance was recorded for all species by 16–23 underwater stationary visual censuses per site (sampling times were not reported). Red hind size and survival data was recorded for a total of 75 individuals from October 2002 to December 2003 by a tagging study totalling 60 fishing events (one/site/month).

    Study and other actions tested
  32. A site comparison study in 2004–2005 at three island coral reef sites in the Indian Ocean, off Tanzania (McClanahan et al. 2009) found that a small marine protected area where all fishing had been prohibited for 13 years had a greater fish biomass compared to areas that have no fishing restrictions. Fish biomass was greater in the area that prohibited fishing (886 kg/ha) than two nearby areas where fishing is allowed (283 and 291 kg/ha). The privately owned Chumbe Island Coral Park off Zanzibar was established in 1991 (0.3 km2, all extractive activities prohibited). In 2004–2005, fish were surveyed by underwater visual census (5 × 100 m belt transects) at two sites inside the protected area and two fished sites with no management 20 km away. Fish >3 cm were recorded by family group and 10 cm size categories, and biomass estimated from length–weight relationships.

    Study and other actions tested
  33. A before-and-after, site comparison study in 1993–2001 of an area of coral reef in the southwest Pacific Ocean, off New Caledonia (Preuss et al. 2009) found that over a nine year period, a marine reserve closed consecutively to all fishing for 5 years had higher overall fish species richness, but not fish abundance and biomass, compared to an area with changing fishing restrictions (initially closed, then opened for two years, then closed again) during the same period. Average fish species richness varied between years but was greater overall in the permanent closure area (permanent: 21–24, non-permanent: 19–21). Overall fish abundance and biomass declined over time in both the permanent (abundance, 1993: 201, 2001: 133 fish/transect; biomass, 1993: 45, 2001: 13 kg/transect) and non-permanent closure areas (abundance, 1993: 220, 2001: 163 fish/transect; biomass, 1993: 27, 2001: 15 kg/transect) but no effect of changes in area management were detected. However, differences between areas were reported for fish species groups divided up by size, feeding habit, mobility, and interest to fisheries (see paper for group individual results). Fish were surveyed at the Aboré reef reserve (15,000 ha, all fishing prohibited in the entire reserve in August 1988) in two areas with different management regime histories: one third closed to fishing since 1998, and two thirds closed in 1988, opened from September 1993 and closed again in September 1995. Diver underwater visual surveys were done in July 1993 (60 transects across entire closure area), July 1995 (48 transects in closed and 62 in open areas) and 2001 (42 transects across entire closure). Transects were 50 m long and fish were recorded by species and size.

    Study and other actions tested
  34. A replicated, site comparison study in 2006–2007 in six areas of seagrass Posidonia oceanica bed in the Mediterranean Sea, France (Seytre & Francour 2009, same experimental set-up as Seytre & Francour 2008) found higher abundance of some fish groups and higher species richness inside a marine protected area in which all fishing was banned compared to two fished reference areas, three years after designation. In May 2007, total biomass, species richness, total fish density and density of four fish families were similar during visual transects over seagrass beds inside and outside the reserve. In September 2007, total biomass, species richness, total fish density and that of two of four families were similar inside and outside the reserve, but density of breams Sparidae spp. and groupers Serranidae spp. were higher inside (bream: 9, grouper: 1/40 m2) than outside the reserve (bream: 5–10, grouper: 0–1/40 m2). Experimental netting data showed higher total abundance, abundance of scorpionfish Scorpaenidae spp. and species richness inside the reserve than outside the reserve in October 2006 (total: 4–9, scorpionfish: 1–3, richness: 3–5/100 m) and June 2007 (total: 1–2, scorpionfish: 0–1, richness: 1–2/100 m). The Cap-Roux Marine Protected Area (450 ha) was closed to all fishing in 2003. Two stations were sampled in each of three zones, one inside the reserve and two outside (north and south) of the reserve boundaries. At each station six replicate underwater visual censuses over seagrass beds and six 100 m trammel net deployments were undertaken. Sampling was undertaken seasonally from 2005–2007. Only data from 2006 were included in the analysis. For full sampling details see original study.

    Study and other actions tested
  35. A site comparison study in 2003–2005 of an area of rocky seabed in the north-western Mediterranean Sea, off Spain (Stelzenmuller et al. 2009) found that fish (functional) diversity and catch rates in local traditional fisheries were greatest closer to a marine reserve closed to all fishing for 22 years, and decreased with increasing distance from the reserve. Functional diversity (the roles played by different species in the ecosystem) and fisher catch rates increased with decreasing distance from the reserve (data reported as statistical model results). However, these were also strongly affected by the presence of a seagrass bed along the western border of the reserve buffer zone. Species diversity was also highest in the waters surrounding the area protected from fishing compared to further away and changed with depth (data reported graphically). In addition, the value of catches were highest within the buffer zone of the reserve. Data were collected between March and December from 2003–2005 in two areas: the buffer zone (418 ha, only artisanal fishing allowed) surrounding the Medes Islands marine reserve (93 ha, designated in 1983, all fishing banned inside the reserve in 1991), and in the adjacent unprotected fished area up to 9 km away from the buffer zone. A total of 44 trammel net fishing operations were sampled and 1,685 fish were counted, identified and fish length recorded.

    Study and other actions tested
  36. A site comparison study in 2004 of the water column around an island in the Mediterranean Sea off Mallorca, Spain (Crec'hriou et al. 2010) found that there were more eggs of four commercially targeted fish species inside a no-take (no fishing) marine reserve enforced for three years and in which the adult fish were more abundant, compared to outside (thus protecting a spawning area from fishing and increasing the likelihood of successful egg production). The eggs of all four species/groups (scorpion fish Scorpaena spp., Mediterranean rainbow wrasse Coris julis, brown meagre Sciaena umbra and grouper Epinephelus spp.) were distributed in higher densities inside the non-fished reserve compared to locations outside, up to two nautical miles away (data reported as statistical results and presented graphically). In addition, there was a clear gradient of decreasing egg density with distance away from the reserve for wrasse and grouper. Plankton was collected in July 2004 inside and outside the marine section of the Cabrera National Park (87 km2, designated 1991, enforced 2001) by two methods: bongo nets (27 stations inside and outside, repeated four times) and fixed nets (9 stations inside). Bongo nets were deployed in down and up oblique tows between the surface and 10 m off the seabed (down) and horizontally for five minutes at 20, 10 and 2 m depths (up). Fish eggs (sub-sampled over 200) were identified and counted in the laboratory under a microscope.

    Study and other actions tested
  37. A replicated, site comparison study in 1992–2006 of nine coral reef sites in the Indian Ocean, Kenya (McClanahan et al. 2010) found that the effects of closing protected areas to all fishing for over 15 years on fish abundance varied between species, compared to adjacent openly fished areas, and the effects on fish species richness and diversity varied with the method used to assess them (fisheries independent underwater visual census versus fisheries dependent catch data). Across bot sampling methods, fish abundances differed between non-fished and fished areas, with about half of the species recorded as common to both management areas by each method being more abundant in closed areas (data reported graphically and as statistical results). Visual census sampling found that the total number of fish species was higher in closed (134) than fished areas (94) and species diversity was similar (0.94–0.95). In contrast, trap and line fishing methods found lower numbers of fish species and diversity at closed areas (number, closed: 30–79, fished: 73–107 species; diversity, closed: 0.5–0.8, fished: 0.8). Trap and line fishing data was collected from two fisheries dependent sources: experimental catch and release studies undertaken for scientific purposes in three old Kenyan fisheries closures (established by 1978, all extractive activities prohibited) and catch composition measured from adjacent traditional fisheries using the same gear types at six heavily fished grounds; and compared with existing fisheries dependent data (underwater visual census surveys, see paper for studies) collected at the same sites. Experimental fishing took place between 1995 and 2006 at three closed areas: Malindi and Watamu Marine National Parks (traps only) and Kisite Marine National Park (traps and line); and six fishing grounds (both methods, line catches obtained from local traditional fishers). Visual census surveys were done at all sites between 1992–2006 (see original paper for details).

    Study and other actions tested
  38. A replicated, site-comparison study in 2005 of shallow rocky reefs in three marine protected areas established for 9–15 years in the Atlantic Ocean around the Canary Islands, Spain (Clemente et al. 2011) found that banning all types of fishing resulted in a different fish assemblage and increased abundance of grey triggerfish Balistes capriscus and barred hogfish Bodianus scrofa compared to unprotected (fished) reference areas, but abundance of ocean triggerfish Canthidermis sufflamen and ornate wrasse Thalassamo parvo was similar. Across all three islands, the fish assemblage was different in unfished and fished areas. Average abundance of grey triggerfish and barred hogfish was higher in the unfished areas (triggerfish: 0.3–0.6; hogfish: 0.1–0.4 fish/100 m2) than the fished areas (triggerfish: 0.0–0.2; hogfish: 0.0 fish/100 m2). Similar abundance in unfished and fished areas was recorded for ocean triggerfish (0.0–0.2 vs 0.1 fish/100m2) and ornate wrasse >12 cm (22.3–35.0 vs 21.7–31.2 fish/100m2). Also reported, but not statistically tested, was abundance of zebra seabream Diplodus cervinus >30 cm (unfished: 0.7-1.3; fished: 0.2–0.6 fish/100m2) and white seabream Diplodus sargus >30 cm (unfished: 0.2–0.7; fished: 0.1–1.0 fish/100m2). Fish surveys were done in April-November 2005 at four sites in each of three marine protected areas and three nearby unprotected fished areas. Protected areas were designated 9–15 years prior, prohibited all fishing. At each site, divers recorded the number and length of all fish for 5 minutes within a 100 m2 circle at six replicate locations.

    Study and other actions tested
  39. A systematic review in 2011 of 32 studies of marine reserves in the Atlantic and Pacific Oceans off Latin America and the Caribbean (Guarderas et al. 2011) found higher total (fish and invertebrates) density, biomass and body size, but not species richness, inside protected areas where fishing is prohibited compared to unprotected fished areas, and the response of fish species was greater than other groups. Data were reported as response ratios. At the food chain level, the increase in density and biomass of fish predators in no fishing reserves was greater than herbivores (fish and invertebrates combined), macroalgae and corals, and at the species level, the increase in density of fish species was greater than invertebrate species. In addition, there was no relationship between the size or age of reserves or area surveyed and the species density (fish and invertebrates) response, but it was found to be associated with one of four variables, intensity of exploitation outside the reserve (i.e. the higher the exploitation level the greater the species response). The systematic review summarized the effects of protection from fishing activities at 23 marine reserves established from 1959 to 2001 in Latin American and Caribbean countries. Four publication databases were searched from 1970–2007 and fifteen site comparison and five before-and-after site comparison studies selected. Data from inside and outside the reserves and before and after designation were used to calculate response ratios to represent the size of the effect of protection.

    Study and other actions tested
  40. A replicated, site comparison study in 2005–2006 of 20 reef sites in the Indian Ocean off Kenya and the Maldives (McClanahan 2011) found that prohibiting all fishing in protected areas (Kenya) for over 25 years resulted in higher total fish biomass, density and species richness compared to heavily fished unprotected areas, but not to partially fished management areas (Maldives). Total biomass in the fully closed areas (1,180 kg/ha) was similar to partially fished areas (1,463 kg/ha), and both were higher than fished areas (110 kg/ha). For nine fish family groups, total density differed between all three areas and was highest at partially fished areas (closed: 463, partly fished: 602, fished: 202 fish/500m2); and species richness was similar at closed and partly fished areas and higher than fished (closed: 45, partly fished: 45, fished: 26 species/500m2). In addition, the effects of different management regimes varied for individual family groups (see paper for data). Fish were surveyed by underwater visual census in four well enforced marine protected areas in Kenya (total 54 km2, established in the 1970s, all fishing prohibited) and four nearby heavily fished areas; and in the Maldives, at 12 sites in a large, lightly managed fished area (650 km2, selectively fished, non-enforced closure system). Sampling took place in February to May 2006 (Kenya) and June 2005 (Maldives). Fish biomass for 23 families was sampled by one or two separate passes along four 5 × 100 m belt transects/site and data for nine selected families by four passes along the 500m2 transects.

    Study and other actions tested
  41. A replicated, site-comparison study in 1996–2009 at eight coral reef sites in the Caribbean Sea, off Belize (McClanahan et al. 2011) found that over a 15 year period following closure of an area of a marine reserve to all fishing, there was a higher number reef fish species and higher abundance of some species groups compared to nearby fished reefs in the reserve, and the effect varied with level in the food chain. The total number of species was higher at unfished reefs (19–27) than fished reefs (17–20) and increased with time. Seven of 17 fish family groups were more abundant (individuals observed/5 min) inside than outside the reserve, nine were similar, and one (Pomacentridae) was more abundant outside (see paper for individual data by group). Snapper abundance (Lutjanidae) showed the largest increase inside the reserve over time (13–72), whilst remaining constant outside (7). Average abundance of carnivorous fish was higher inside than outside the reserve, including: fish-eating fish (16 vs 4); fish that feed on large invertebrates (2 vs 1) and fish that feed on small invertebrates (159 vs 126). Abundance of herbivorous fish (284 vs 298) and sponge-eating fish (1 vs 1) was similar inside and outside the reserve. Fish were surveyed by underwater visual censuses at four reefs in the conservation zone (71 km2, legal protection in 1993, no fishing since 1995) and four nearby reefs in the general use zone (190 km2, regulated fishing activity) of Glover’s Reef Marine Reserve. Each reef site was sampled 8–10 times during May-November between 1996–2009. Divers haphazardly swam over each reef for a total of 35 minutes and recorded the number and species of fish from seven taxonomic groups during separate 5-minute intervals.

    Study and other actions tested
  42. A replicated, paired, site comparison study in 2009–2010 of four estuaries in the Tasman Sea, New South Wales, Australia (McKinley et al. 2011) found higher average abundance of commercially targeted fish and similar species number in marine park zones where all fishing has been prohibited for four to eight years, compared to partially fished park zones, but there was a lower abundance of targeted fish and a different overall fish assemblage than unprotected fished estuaries. Abundance in no fishing park zones was higher than fished park zones for all targeted fish (no fishing: 3.9, fished: 1.5 count/camera drop), and individually for pink snapper Pagrus auratus (no fishing: 0.9, fished: 0.1 count/camera drop), but was similar for silver trevally Pseudocaranx georgianus and yellowfin bream Acanthopagrus australis (trevally: 1.0 vs 0.0, bream: 0.2 vs 0.1). The number of fish species (no fishing: 6.3, fished: 4.8) and maximum abundance of all fish (no fishing: 27, fished: 45 fish/camera drop) were similar in non-fished and fished park zones. In addition, targeted fish abundance was higher in estuaries without marine parks (12 fish/camera drop) and had a different fish assemblage (data reported graphically). Four estuaries, in New South Wales (100–400 km apart) were randomly sampled from November 2009 to March 2010 using baited remote underwater video. Two estuaries were marine parks (four and eight years old), zoned into no fishing areas and areas where some commercial and recreational fishing (netting and trapping) was permitted. The other two estuaries had no conservation designation and although most commercial fishing was banned they were intensively fished recreationally.

    Study and other actions tested
  43. A replicated, site comparison study in 2005 of seven coral reef areas in the Indian Ocean off the coasts of South Africa and Mozambique (Currie et al. 2012) found that six years after prohibiting all fishing in no-take areas of marine reserves there was increased abundance of six of 12 fish species/groups compared to partly fished and openly fished areas, and overall fish size was larger than in openly fished areas. Average abundances were higher inside no-take areas than partly fished and openly fished areas for six of 12 fish species/groups (no-take: 0.5–9.0 fish/count, part fished: 0.1–3.0 fish/count, open: 0.0–3.0 fish/count). The abundances of the other six were higher in no-take areas compared to partly fished but were similar to openly fished areas (no-take: 0.3–10.0 fish/count, part fished: 0.0–6.0 fish/count, open: 0.3–10.0 fish/count). See original paper for list of species and individual abundances. Average fish size (reported as a standardised measure) across the whole assemblage was higher inside no-take areas (57) than openly fished areas (48) and similar to partly fished areas (58). In April 2005, fish were sampled at two no-take areas (no extractive activity) and four partly protected areas (limited non-commercial/non-trawl fishing types and diving permitted) in adjacent marine reserves (designated 1999), and at five openly fished sites outside the reserves (two adjacent and three >200 km away). At each site, divers counted selected fish species >7 cm in length, along two replicates of bisecting transect pairs 25 m long and 5m wide. Point counts (22–32) were also conducted at each site in a 5 m radius, separated by 20 m. Data were analysed for seven coral-dominated sites (two no-take and open, three part protected).

    Study and other actions tested
  44. A replicated, paired, site comparison study in 2004–2010 of five coral reef regions in the Great Barrier Reef Marine Reserve, Australia (Miller et al. 2012) found that reefs closed to fishing for two to six years had greater numbers of coral trout Plectropomus/Variola spp., compared to fished reefs. Across all years and reef regions, the total number of coral trout was greater at reefs closed to fishing (0.8–0.9 fish/tow) than open reefs (0.4–0.6 fish/tow). Similarly, overall coral trout number at each of the five individual reef regions was higher at sites closed to fishing (Cairns: 0.2–0.3 fish/tow; Townsville: 0.5–0.7; Mackay: 1–1.1; Swains: 1.4–2.6; Capricorn Bunker: 0.5–0.8) compared to their paired, fished sites (Cairns: 0.1–0.2 fish/tow; Townsville: 0.1–0.2; Mackay: 0.1–0.5; Swains: 0.2–1.3; Capricorn Bunker: 0.2–0.4). In 2004, the Great Barrier Reef was rezoned to create no–take marine reserves. In 2006–2010, a total of 28 pairs of reefs were surveyed across five reef regions, 25–450 km apart (six pairs in each region except Capricorn Bunker, where four pairs were surveyed). Each reef pair was one reef closed to fishing and one fished (0-1 km apart). Fish at each reef site were sampled by the manta tow method, where 10 m-wide areas of reef slope are surveyed at a time by an observer towed behind a small boat, for two minutes. Paired sites were surveyed within 12 months of each other on a biennial basis over six years.

    Study and other actions tested
  45. A site comparison study in 2009–2010 at seven mangrove and coral reef sites in Moreton Bay, Coral Sea, Australia (Olds et al. 2012) found that prohibiting all fishing inside a marine reserve for over 12 years resulted in greater fish density of three of four fish groups at coral reefs and one of four in mangroves, compared to non-reserve areas, and was influenced by proximity to other habitat types. At no-take coral reef areas close to mangroves, fish density of three of four fish groups was higher than non-reserve areas (harvested: 65–159 vs 42–96, herbivorous: 75–138 vs 34–79, piscivorous: 22–39 vs 13–26 fish/200 m2), but the density of prey fish species was lower (reserve: 77, non-reserve: 145 fish/200 m2). Reserve coral reef areas far from mangroves had greater fish density for piscivorous fish only compared to non-reserve areas (28–30 vs 19–22 fish/200 m2). Mangroves in reserve areas near coral reefs had greater densities of piscivorous fish (reserve: 37, non-reserve: 18 fish/200 m2) but lower densities of prey fish (reserve: 41, non-reserve: 255 fish/200 m2). Reef fish were surveyed in summer 2009–2010 inside a no-take marine reserve, protected since 1997, and at six non-reserve sites in Morton Bay (0–25 km away). At each site, two coral and three mangrove areas were sampled. On coral reef, fish were sampled along five, 50 by 4 m transects at each site by underwater visual census. Fish in mangroves were surveyed at high tide using underwater transects and fyke nets.

    Study and other actions tested
  46. A replicated, site-comparison study in 2009–2011 at five coral reefs in the Coral Sea, Australia (Olds et al. 2012) found that prohibiting all fishing within a no-take marine reserve for over 12 years increased the diversity and biomass of herbivorous fish compared to non-reserve reference areas at reefs close to, but not distant from, mangrove forests. At reefs close to mangroves, herbivore species richness was higher inside the reserve (8 species/200 m2) than outside (5 species/200 m2), but similar for reefs further away from mangroves (inside: 5, outside: 4 species/200 m2). Herbivore biomass at reefs close to mangroves was also higher inside the reserve (inside: 14, outside: 7 g/m2), mainly due to the higher biomass of roving browsers and black rabbitfish Siganus fuscescens (data reported as statistical model results), and similar at distant reefs (inside: 3, outside: 2 g/m2). In addition, across both near and far reefs the biomass of roving grazers, the Australian sawtail Prionirus microlepidotus and the blue-barred parrotfish Scarus ghobban was higher at reserve than non-reserve reefs (data reported as statistical model results). Fish were surveyed along five replicate 50 × 4 m underwater transects at low tide at one protected reef and four unprotected reefs in the Moreton Bay Marine Park, eastern Australia, from November 2009 to January 2011. The protected reef is a no-take reserve where all fishing is banned (since 1997). At each location one reef close (<250 m) to mangroves and one distant (>500 m) from mangroves were sampled.

    Study and other actions tested
  47. A replicated, paired, site comparison study (year not stated) of three marine reserves in the Mediterranean Sea, Spain (Verges et al. 2012) found that prohibiting all fishing resulted in higher biomass of predatory, but similar biomass of herbivorous, fish inside one no-take marine reserve compared to outside unprotected areas, and similar biomass to two other marine reserves with a different level of protection from fishing. The total biomass of predatory fish was higher inside than outside (inside: 32,522, outside: 13,984 g/250m2) at the only no-take reserve (Catalunya) and was similar at the two other reserves (inside: 10,025–15,699, outside: 6,484–18,815 g/250m2). No effect of protection level was found on the total biomass of herbivorous fish at all three reserves (inside: 5,322–15,000, outside: 3,064–4,516), but it was influenced by an interaction of protection, depth and reserve region (data reported as statistical results). At each reserve, three sites were sampled inside and three outside the reserve boundaries, one at each depth of 5, 15 and 30 m (date or year of sampling unspecified). Fish were identified and counted by diver underwater visual survey along three, 50 × 5 m transects at each site. Herbivorous fish and fish that predate on sea urchins Paracentrotus lividus were recorded (see paper for list of families). The Catalunya reserve was protected since 1983 and prohibits all extractive activities. The other two reserves were protected since 1991 (Mallorca) and 1999 (Menorca) and permit some restricted commercial fishing.

    Study and other actions tested
  48. A replicated, paired, site comparison study in 2008 of five shallow rocky seabed areas in the Mediterranean Sea, Spain (Villamor & Becerro 2021) found that no-take marine reserves closed to fishing for at least 10 years, had higher overall abundances of top predators (fish) and carnivore species (fish and invertebrates) and a similar abundance of herbivore species (fish and invertebrates), compared to non-protected areas outside. Overall, top fish predators (two families) and carnivores (three fish families, one invertebrate) were more abundant inside than outside the no-take marine reserves and the abundance of herbivores (one fish and one invertebrate species) was similar (data reported as statistical results and presented graphically for each reserve). In August 2008, five marine reserves along the east coast of Spain were surveyed by underwater visual census. Six transects, 50 × 5 m, were done at each reserve: three in no-take areas and three in unprotected areas nearby (4–12 km). All fish, and two invertebrate species, were identified, counted, and assigned to one of three universal feeding groups (see paper for list of species). Reserves were protected for 10–25 years.

    Study and other actions tested
  49. A replicated, site comparison study in 2010–2011 at six coral reef sites in a marine park in the Indian Ocean, Western Australia (Wilson et al. 2012) found that the level of protection from fishing did not influence fish abundance, biomass and diversity between zones where no fishing was permitted for five to 20 years and fished zones. Total fish abundance, biomass and diversity of adult fish was similar between unfished and fished zones (data presented as fitted model outputs and statistical results), but some differences were found for fish grouped together based on diet/feeding behaviour (see paper for results by fish group). Sanctuary zones (free from fishing) in Ningaloo Marine Park were established in 1991 and 2005. At each of six sites within the reserve; three where no fishing is allowed, and three where some commercial fishing is permitted, a total of 9–14 patch reefs 2–4 m deep were surveyed. All adult fish visible on each reef were identified and counted by a single underwater observer in November 2010–January 2011.

    Study and other actions tested
  50. A before-and-after, site comparison study in 1996–2005 of three seabed areas on the Patagonian Shelf, South Atlantic Ocean, off Argentina (Alemany et al. 2013) found a different assemblage of bottom dwelling fish, higher overall abundance and higher abundance of target Argentinian hake Merluccius hubbsi, and particularly of young hake, in a marine protected area in which fishing was banned for up to eight years, compared to two fished reference areas. The whole fish assemblage before the closure was similar in the protected area to one of the two outside areas but differed from both outside areas after the closure (data reported as statistical model results). Before the closure, overall fish abundance was similar inside (0.59 t) and outside (0.45–0.79 t) the reserve, but increased inside the reserve following the closure and was higher than outside in two of four years (inside: 0.73–0.88, outside: 0.27–0.54 t). Hake abundance was similar across areas before the closure (inside: 0.54, outside: 0.26–64 t) but increased inside the reserve and was higher after the closure in all years (0.52–0.89 t) relative to outside (0.13–0.61 t). The proportion of two-year old hake inside the reserve was higher after the closure (36–50%) than before (18%). The Patagonian Closed Area (50–100 m depth) was closed to all fishing in 1997. Data from demersal fish surveys (5 × 30 m trawl with a 2.4 cm codend mesh) before (1996) and after (2000–2003, 2005) the closure were analysed from a selected 28,000 km2 area inside the reserve and two fished areas outside. All fish were counted, identified, and the ages and lengths of hake recorded.

    Study and other actions tested
  51. A replicated, before-and-after, site comparison study in 1999–2011 at four coral reef sites in the Gulf of Mexico, off Florida Keys, USA (Ault et al. 2013) found that in marine reserve areas where all fishing was prohibited for up to 10 years, and in areas where only recreational fishing is permitted, there were increases in the density of commercial fish species in the 10 years following implementation and compared to openly fished areas, and changes in fish densities of non-target and other exploited fish species varied. For five fishery exploited species, the total number of increases in density detected in surveys was higher overall in no-take and recreationally fished areas compared to openly fished areas (no-take: 3, recreational: 16, fished: 1), and decreases were only detected in the openly fished areas (no-take: 0, recreational: 0, fished: 5). For non-target species (increase, no-take: 9, recreational: 20, fished: 12; decrease, no-take: 9, recreational: 12, fished: 3) and species collected for the aquaria trade (increase, no-take: 0, recreational: 7, fished: 1; decrease, no-take: 9, recreational: 7, fished: 3), changes in density fluctuated between years (see paper for species individual data). Fish were surveyed over 326 km2 at four sites with three different levels of resource management protection; Tortugas North and South Ecological Reserves (no-take, since 2001), Dry Tortugas National Park (part no-take, since 2007 and part recreational angling only, since 1960s in all areas). Baseline fish surveys were done in 1999–2000 before the no-take areas were implemented and from 2002–2011. Diver visual surveys were done in a two-stage stratified random sampling design. Numbers of reef fish were recorded in randomly selected circular plots 15 m in diameter.

    Study and other actions tested
  52. A site comparison study in 2009–2010 of 12 rocky reef and boulder sites in the Mediterranean Sea, off eastern Sicily, Italy (Pierpaolo et al. 2013) found that five to six years after all fishing was banned in a reserve zone of a marine protected area, the overall fish assemblage was different and fish abundance, species richness and diversity was higher compared to fished areas outside the reserve, but the effect on individual species abundance varied between size classes and commercial/non-commercial species. The overall fish assemblage was different inside and outside the reserve (data reported as statistical results) and total fish abundance was higher inside (226 fish/125 m2) than outside (90 fish/125 m2). This was due to greater abundances of medium and large fish inside (medium: 80, large: 108 fish/125 m2) than outside (medium: 38, large: 25/125 m2), as well as species of high commercial value (24 vs 4 fish/125 m2). Abundance of small and low and medium commercial value fish were not significantly different inside and outside the reserve (small: 38 vs 27 fish/125 m2, low value: 12 vs 8 fish/125 m2, medium value 67 vs 4 fish/125 m2). Fish species richness and Shannon diversity were higher inside the reserve (species: 14, diversity: 1.7 fish/125 m2) than outside (species: 12, diversity: 1.5 fish/125 m2). Fish were surveyed by underwater visual census in early summer 2009–2010 along three 125 m2 transects (15–20 m depth) at four sites inside the marine reserve zone (where all fishing activities are prohibited since 2004) of the Plemmirio Marine Protected Area (2,400 ha), and eight sites outside the reserve (four in an adjacent zone where only some controlled fishing activities are allowed and four outside the marine protected area, 12 km away).

    Study and other actions tested
  53. A site comparison study in 2006–2008 of a shallow, sandy lagoon in a bay in the South Atlantic Ocean, off South Africa (da Silva et al. 2013) found that common smoothhound sharks Mustelus mustelus spent more time in a no-take marine protected area than outside (and thus more protected from fishing), and movements between the areas differed with season. Overall, sharks spent an average of 74–80% of hours inside the no-take area over a two-year period. The highest numbers of detections inside the no-take area occurred in summer and the lowest in winter (data presented graphically and as statistical results). In November 2006, a total of 24 smoothhound sharks were tagged with acoustic transmitters and released in the Langebaan Lagoon Marine Protected Area (34 km2, year implemented not reported), a no-take area in the innermost part of a coastal embayment (Saldanha Bay). The movements of sixteen sharks (9 females, 7 males) detected for at least one year, and of nine detected for two years, were analysed. Fish movement detection data was recorded by 28 acoustic receivers positioned at four sites in no-take and fished areas.

    Study and other actions tested
  54. A replicated, paired, site comparison study in 2004–2005 of coral reef and seagrass sites at three neighbouring inhabited islands in the Java Sea, Indonesia (Campbell et al. 2014) found that prohibiting fishing in areas of a 16 year old national park resulted in similar individual fish size and weight in catches landed at two of the three islands, compared to fished areas. Average fish length and weight in landed catches were similar between closed and fished areas at the islands of Karimunjawa (length, closed: 235 mm, fished: 222 mm; weight, closed: 482 g, fished: 395 g) and Parang (length, closed: 317 mm, fished: 311 mm; weight, closed: 733 g, fished: 766 g) and were lower in closed areas at Nyamuk island (length, closed: 306 mm, open: 411 mm; weight, closed: 781 g, open: 1,040 g). Karimunjawa National Park (111,625 ha) was first legislated in 1988 and has zones prohibiting fishing and designated fishing zones. In January 2004–December 2005, fish catch surveys were done by trained observers at 1–2 month intervals at fish landing sites on Karimunjawa Island. A total of 8,674 fish from 895 fishing trips were sampled. Fishers were asked to provide details of where they were fishing and the location of fish capture was assigned to one of five village fishing grounds on separate islands 6–15 km apart. Fishing was reported from both closed and fished management zones off three islands and the fish data compared.

    Study and other actions tested
  55. A replicated, paired, site comparison study in 2012 of six coral reef sites in a marine protected area in the Coral Sea, Vanuatu (Januchowski-Hartley et al. 2014) found that permanent no-take reserves where fishing was prohibited for at least six years had greater total fish biomass compared to areas open to fishing, and similar fish biomass to closed areas fished only periodically for short periods. The total fish biomass was similar in no-take reserves (646–835 kg/ha) and periodically fished areas (559–567 kg/ha) but was greater than fished areas (331–378 kg/ha). The biomass and abundance (data not reported) of only one of three individual fish groups differed between areas and was higher in no-take reserves than the other two areas (see original paper for individual data). Data was collected in November–December 2012 in two regions of the Nguna-Pele Marine Protected Area Network. Each region had three adjacent management zones (8 to 16 ha) that were each surveyed: a no-take reserve (since 2005), a periodically fished area open for 1–3 days every 6 months (implemented since 18 months to 6 years) and an area open to fishing. At each zone before and after a three-day harvesting period of the periodically fished zone, divers recorded fish species and length along eight, 50 by 5 m transects, and biomass calculated from length-weight relationships.

    Study and other actions tested
  56. A site comparison study in 2005–2010 of three coral reef areas in the Caribbean Sea off Puerto Rico (Mateos-Molina et al. 2014) found that prohibiting all fishing in a marine protected area resulted in a similar coral reef fish abundance and biomass one and five years after implementation compared to fished areas, but abundance increased in all areas over time. Overall, there were no differences in average reef fish abundance and biomass between no-take and fished locations, but after 5 years abundance had increased in all areas, regardless of protection level, particularly for small life stages and small-sized fish (data presented graphically and as statistical results). A no-take zone at the Mona Island Marine Protected Area was established in 2004 extending up to 926 m from the shore initially and modified in 2007 to include areas up to 182 m depth. Two locations in the no-take area and one in a fished area of the marine protected area (within the 2004 boundaries) were surveyed in autumn and winter of 2005–2006 and 2009–2010. At each location, fish size and abundance were recorded by underwater visual census along 12 belt transects (60 m2) at three separate sampling sites. After each transect five-minute roving surveys were conducted.

    Study and other actions tested
  57. A replicated, paired, site comparison study in 2011 of six mixed reef, mangrove and seagrass lagoon areas in the Soloman Sea, Soloman Islands (Olds et al. 2014) found that no-take marine reserves protected for eight years had higher fish abundances than unprotected fished sites for four of six species, but the effect differed with type and proximity of different habitats. Fish density of four of six species was higher in at least two of the five habitat categories in no-take reserves compared to fished areas (bumphead parrotfish Bolbometopon muricatum: 2–6 vs 0, mangrove snapper Lutjanus argentimaculatus: 4–5 vs 0–1, goldlined rabbitfish Siganus lineatus: 5–31 vs 0–5, ringtail surgeonfish Acanthurus blochii: 5–15 vs 1–3 fish/200 m2). For two species, density was similar between areas in four of the habitats and was lower in reserves in one (monocle bream Scolopsis spp: 5 vs 8, dash-and-dot goatfish Parapeneus barberinus: 1 vs 7 fish/200 m2). In addition, the authors reported increases in abundance in reserves of a total of 18 fish species (data presented in the Supporting Information). Three small, community-based no-take reserves (established eight years) designed for bumphead parrotfish, and three paired unprotected fished locations were surveyed in April-June 2011. At each location, fish over 5 cm length were recorded by underwater visual census (5 × 200 m2 transects) in mangrove, seagrass and coral reef habitats. Fish data were assigned to one of five categories: mangroves near coral, coral near mangroves, isolated coral, coral near seagrass and seagrass near coral.

    Study and other actions tested
  58. A site comparison study in 2010–2013 of a fished area of seabed in the north Atlantic Ocean off the Isle of Arran, Scotland, UK (Howarth et al. 2015) found that prohibiting all types of fishing in a marine reserve resulted in similar overall fish abundance, similar abundances of seven of seven individual fish groups, and similar sizes of four of four fish groups compared to an adjacent fished area outside the reserve, up to five years after implementation. Across years, overall fish abundance (total number) was not statistically different between non-fished reserve and fished areas (reserve: 803, fished: 644) and the maximum numbers of seven of seven fish groups, dominated by cod Gadus morhua and other ‘cod-like’ fish Gadidae, were similar (reserve: <1–9, fished: <1–9; see paper for individual data by fish group). Fish size was similar between the reserve and fished areas for cod, other cod-like fish, flatfish Pleuronectidae and lesser-spotted dogfish Scyliorhinus canicula (data reported as statistical results; three fish groups were not tested). Lamlash Bay Marine Reserve (2.7 km2) was established in September 2008 and closed to all fishing. Annually between 2010–2013, fourteen to 20 sites inside and outside the reserve were sampled. Fish data were collected by diver visual surveys along 150 m2 transects (total number) and analysis of footage recorded by baited remote underwater video (species, number and fish length).

    Study and other actions tested
  59. A replicated, site comparison study in 2009–2011 of 23 coral reef sites spanning four regions in the Pacific Ocean (Phillipines, Papua New Guinea, Vanuatu) and Indian Ocean (Chagos) (Januchowski-Hartley et al. 2015) found that surgeonfish and parrotfish inside established marine protected areas where fishing was banned showed the same pattern of increasing avoidance behaviour (measured as flight initiation distance) with increasing fishing intensity in the locality, compared to fish in fished areas. Flight initiation distance increased in both non-fished and fished sites with increasing local fishing pressure levels (lowest to highest) for surgeonfish Acanthuridae spp. (from 155 to 222 cm in non-fished areas and 270 to 408 cm in fished) and parrotfish Scaridae spp. (from 211 to 279 cm in non-fished areas and 332 to 537 cm in fished). In 2009–2011 thirteen sites protected from fishing through permanent no-take reserves or traditional management closures (reserve size or year of implementation were not reported) and 10 sites that allowed fishing were surveyed across four countries. Fish flight initiation distance was estimated by measuring how closely a diver could approach individual fish (> 10 cm total length) before they fled. Fishing pressure was estimated by dividing the linear extent of reef open to fishing by the number of fishers in the fishing community and ranged from 0–80 fishers/km.

    Study and other actions tested
  60. A replicated study in 2001–2013 of four surf-zone sites in the Indian Ocean, off South Africa (Mann et al. 2015) found that over a nine-year period, the majority of recaptures made of tagged fish from five species, occurred inside a marine reserve where fishing activity was controlled by zones, and mainly within 200 m of their original release site in the no-take reserve zone closed to all fishing for over 22 years (and thus spent more time in areas protected from fishing). Most individuals of the five main study species were recaptured within 200 m of their original release site (grey grunter Pomadasys furcatus: 88%, catface rockcod Epinephelus andersoni: 84%, yellowbelly rockcod Epinephelus marginatus: 92%, cave bass Dinoperca petersi: 88% and speckled snapper Lutjanus rivulatus: 79%) and 61% of fish were originally tagged at sites in the no-take zone, the rest in the zone that allows shore angling and recreational boat angling and spearfishing for pelagic gamefish only. In addition, the maximum time at liberty of each species ranged from 287–3,163 days, average recapture rate was 29% and 632 of the 3,224 fish tagged were recaptured at least once. The St Lucia Marine Reserve in South Africa was established in 1979. From November 2001–2013, a total of 6,613 fish from 71 species were tagged and released at four sites in the reserve: two in a no-take zone and two in a restricted fishing zone. Over the same sampling period, details of fish recaptured in the reserve by the research team and angling public, and other reported recaptures in fished areas outside the reserve were recorded.

    Study and other actions tested
  61. A replicated, site comparison study in 2005–2012 of 233 coral reef sites across the western Indian Ocean (multiple countries) (McClanhan et al. 2015) found that the biomass of the fishable portion of reef fish communities (standing stock biomass) increased across a gradient of decreasing fishing intensity resulting from six different management regimes, and was highest in protected areas closed to fishing and with enforcement. Data were not statistically tested. Average fishable biomass was greatest in large, remote marine protected areas (2,189 kg/ha, 36 sites) and areas closed to fishing with high compliance (957 kg/ha, 114 sites), whereas young areas closed to fishing with low compliance had 489 kg/ha (66 sites). Areas where all (line and traps only) or most (spear and gill nets also used) destructive gears were restricted had 390 and 382 kg/ha of fishable biomass, respectively. The lowest biomass was in areas with no gear restrictions (269 kg/ha, 50 sites, seines and explosives used). In addition, many of the individual sites, even in areas with closures and high compliance, had a fishable biomass below 1,150 kg/ha (estimated by the authors as the target standing stock biomass needed for the recovery of exploited reef fish), and were thus failing to achieve conservation targets. Coral reef fish assemblages were surveyed at 233 individual sites across the Indian Ocean (off Comoros, Kenya, Madagascar, Mayotte, Mozambique, the Maldives, Seychelles, the Chagos archipelago and Tanzania) between 2005–2012. Fish were surveyed at each site by underwater visual census (3 to 5 belt transects of 50 or 100 m, or 8 point counts – see original paper for sampling methods by country). Sites were classified by the six dominant management categories.

    Study and other actions tested
  62. A replicated, site comparison study in 2012–2013 of 37 coral reef sites with at least one established, locally-managed marine protected area in the Philippine Sea, Philippines (Muallil et al. 2015) found that areas where all fishing was prohibited had greater fish species richness and diversity, fish density and larger fish for five out of seven family groups, compared to nearby fished areas. Overall fish species richness and diversity (data reported as diversity indices) was higher in protected areas (20 species) than fished areas (15 species). Density was higher for five of seven reef fish families (surgeonfishes Acanthuridae: 18 vs 16, parrotfishes Scaridae: 9 vs 6, snappers Lutjanidae: 7 vs 6, groupers Epinephelinae: 3 vs 2, goatfishes Mullidae: 2 vs 1 fish/500 m2) and similar for grunts Haemulidae and emperorfish Lethrinidae (both <1 fish/500m2 in all areas). A greater number of larger (25 cm and above) individuals of five families were found at protected sites compared to fished sites (surgeonfishes: 0.8 vs 0.1, parrotfishes: 1.4 vs 0.4, groupers: 0.4 vs 0.2, goatfishes: 0.2 vs 0.1, grunts: 0.13 vs 0.07 fish/500 m2) and similar for snappers (0.7 vs 0.6 fish/500m2) and emperors (0.0 vs 0.0 fish/500m2). Between 2012–2013, reef fish were surveyed at 37 locations by underwater visual census along 348 belt transects (50 × 10 m). At each location, 8–12 transects were done, half in and half outside (>200 m) protected areas. Species, number, and estimated length was recorded for fish above 5 cm. The marine protected areas were mostly <50 ha, and the years since implementation were not reported.

    Study and other actions tested
  63. A site comparison study in 2011–2013 of 18 coral reef sites on the Great Barrier Reef in the Coral Sea, Australia (Rizzari et al. 2015) found that in a marine protected area where human activity was controlled by zones, of six different fish trophic groups, two were more abundant and two had a larger size and biomass in no-entry zones than no-take and fished zones, after 10 to >20 years of protection. Densities of apex predators and browser herbivores were higher in the no-take zone compared to both the no-take and fished zones but there were no differences between areas for targeted and non-targeted medium-sized predators and two other groups of herbivorous fish (data reported graphically and as statistical results). Fish size and biomass differed between areas only for the targeted and non-targeted predator groups and were higher in the no-entry zone than the other zones (data reported as statistical results). In addition, the differences in the predator groups due to protection level were not found to influence the density, size or biomass of the herbivorous fish groups. Reefs in three management zones within the Great Barrier Reef Marine Park were surveyed from October-April 2011–2013: no-entry (protection >20 years), no-take (protected 10–20 years where fishing is prohibited but non-extractive activities like diving are allowed), and fished areas. Fish were categorized into six groups according to food chain position and exploitation status (see original paper for details). At each reef (six per zone), apex predators were surveyed two to six times using 45-minute timed swims (20 m wide transect) and medium-sized predators and herbivores >10 cm total length using 10 to 16 transects (10 × 50 m).

    Study and other actions tested
  64. A replicated study in 1993–2013 of 17 reef areas off Cocos Island, in the eastern Pacific Ocean, Costa Rica (White et al. 2015) found that over a period of 21 years, eight of twelve shark and ray species declined in abundance or presence inside a no-fishing marine protected area established for over 15 years, and poor enforcement may have contributed to the decline. Percentage declines in observed abundance in the period from 1993 to 2013 were recorded for six of twelve species (scalloped hammerhead Sphyrna lewini: 45%, whitetip reef shark Triaenodon obesus: 77%, marble ray Taeniura meyeni: 73%, eagle ray Aetobatus narinari: 34%, mobula ray Mobula spp.: 78%, manta ray Manta birostris: 89%) and declines in the likelihood of occurrence recorded for two (silky shark Carcharhinus falciformis: 91%, silvertip shark Carcharhinus albimarginatus: 87%). The likelihood of occurrence of four species increased between 1993 and 2013: tiger sharks Galeocerdo cuvier (79%/yr, Galapagos sharks Carcharhinus galapagensis (33%/yr), blacktip reef sharks Carcharhinus limbatus (9%/year) and whale sharks Rhincodon typus (5%/yr). In addition, the authors reported inadequate enforcement of fishing controls were likely to contribute to the declines. The Cocos Island National Park was designated in 1978, and extended in 1984 and 2001, covering 22.2 km around the island. Fishing is banned within the park but enforcement is poor and illegal shark fishing occurs. From 1993 to 2013 divers surveyed sharks and rays at 17 sites within the reserve by underwater visual census for one hour. Sites were between 10 and 40 m depth. Common species were recorded as count data and analysed as relative abundance while presence-absence data were recorded for rare species and analysed as odds of occurrence.

    Study and other actions tested
  65. A site comparison study in 2009–2014 of a coral reef area off San Miguel Island in the Philippine Sea, Philippines (Bobiles et al. 2016) found more fish species and a higher overall fish abundance of commercially important fish in a no-entry/no-fishing zone of a marine protected area, compared to two partially fished zones and unprotected fished areas 10 to 15 years after implementation, and the effect of protection varied between individual species groups and sizes. Across all years, the average species richness and fish abundance of commercially important species was higher inside the no-entry zone (species: 11–12, abundance: 28–41 fish/transect) than elsewhere and was similar between partially fished protected zones (species: 3–8, abundance: 5–30 fish/transect) and non-protected fished areas (species: 4–7, abundance: 10–15 fish/transect). For the top six commercial fish family groups, the abundance of market-sized individuals of five groups differed between areas, whereas for non-target sizes only one differed (see paper for individual data). The San Miguel Island Marine Protected Area was designated in 1998 and has three zones with different levels of protection: a 1.0 km2 sanctuary area (no fishing or recreational activity), a 1.25 km2 partially protected area (traditional fishing types - gillnet, spear, trap, longline - permitted), and an outer 100 m buffer protected zone with less restriction (not specified). In May 2009 and 2010 and December 2014, fish were surveyed in each of the three zones and the adjacent unprotected area by underwater visual census along a total of 10 haphazardly placed transects (50 m2) at least 10 m apart. Transects were located at reefs 1.3 km offshore and at depths of 9–21 m.

    Study and other actions tested
  66. A site comparison study in 2013 of a rocky seabed area in the Atlantic Ocean, off southwest Portugal (Gil Fernandez et al. 2016) found that a marine protected area where all fishing activity (except barnacle extraction) is prohibited, had a similar total fish species richness, a higher biomass and size, but not density, of seabream Diplodus spp. and a similar abundance, size and biomass of dreamfish Sarpa salpa and wrasses Labrus and Coris spp. compared to an adjacent fished area after two years. Average fish species richness was similar inside (7.8) and outside (4.5) the protected area. Biomass and size of Diplodus spp. was higher inside (biomass: 262 vs 105 g/100 m2; size: 11.3 vs 5.1 cm), but density was similar (3.9 vs 5.4 fish/100 m2). Similar density, size and biomass were recorded inside and outside the protected area for dreamfish (data not reported), and the wrasses Labrus spp. (density: 0.8 vs 0.3 fish/100 m2; size: 18.4 vs 16.0 cm; biomass: 207 vs 21 g/100 m2) and Coris sp. (density: 2.0 vs 4.5 fish/100 m2; size: 6.4 vs 4.3 cm; biomass: 47 vs 107 g/100 m2). In 2011, all fishing activity bar the extraction of barnacles was banned in the marine section of the Natural Park of the Southwest Alentejo and Vicente Coast (28,858 ha). In February and May 2013, fish were surveyed at two locations inside the protected area and two in an adjacent fished area (all fishing permitted, except bottom trawling and recreational fishing on Wednesdays). At each location two 40 m transects were swum by divers and the number and total length of all fish except small benthic species were recorded.

    Study and other actions tested
  67. A site comparison study in 2012–2014 in shallow, sandy inshore areas in the western Atlantic Ocean off South Caicos in the Turks and Caicos Islands, UK (Henderson et al. 2016) found that banning all fishing in a marine reserve resulted in a higher abundance of immature lemon sharks Negaprion brevirostris, particularly of smaller sizes, and similar shark growth rates but lower condition, compared to fished areas outside, after 20 years. Average abundance of immature lemon sharks was higher inside (0.56 sharks/h) than outside (0.36 sharks/h) the reserve and there were more smaller individuals (data presented graphically). Average condition factor was lower inside the reserve than outside, but growth rates were similar (data reported as statistical tests). The Bell Sound Nature Reserve was established in 1992 to protect bonefish Albula Vulpes and no fishing activity is permitted. Between February 2012 and August 2014, sharks were sampled year-round at 12 sites on a rotating basis (seven inside and five outside) using square-mesh gillnets (100 m long by 1.83 m deep, 6.35 cm mesh size). Nets were set perpendicular to shore for 1–6 h. Sharks (new and recaptured tagged individuals) were removed from nets immediately after capture, weighed and the total length measured. New individuals were marked with both a plastic tag and a data recording tag.

    Study and other actions tested
  68. A site comparison study in 2013–2014 of two coral reef sites in the Indian Ocean off southwest Madagascar (Jaonalison et al. 2016) found that overall species richness and abundance of post-larval fish was similar at reefs where all fishing was prohibited for 15 years compared to fished reefs, and individual fish species or family groups differed with changes in water temperature, salinity and/or transparency. Overall, the non-fished reef had a similar average number of fish species/families (non-fished: 3–6, fished: 1–9) and post-larval abundance as the fished reef (non-fished: 5–24; fished: 2–26). In addition, the most dominant and frequent species/families differed between reefs (see paper for individual data) but this was influenced by sea surface temperature, salinity and water transparency. Two differing reef sites 50 km apart were surveyed monthly (except November) in August 2013–February 2014; a protected reef off Anakao (10 km2, protected from fishing since 1999) and a fished site in the Great Reef of Toliara, with reduced coral diversity. Fish post-larvae were sampled at three locations per site using light-traps and transferred live to a laboratory for identification to the lowest taxonomic level possible.

    Study and other actions tested
  69. A replicated, site comparison study in 2002–2012 of sixteen rocky coral reef sites in a marine park in the Tasman Sea, New South Wales, Australia (Malcolm et al. 2016) found that banning all fishing in areas of the park resulted in increased fish abundances of six of 12 fishery targeted and non-targeted species or groups compared to areas where some fishing types are allowed, and the effect varied with size and age (small, 10–20 years and large, 0–10 years) of the area protected. Across all years, average abundances of five of ten targeted and one of two non-targeted fish species/groups (see paper for details) were higher at a non-fished area than at fished areas, and no effect of management type was found for the other six (data reported as statistical results and presented graphically for some species only). In addition, the effect of management type was generally higher for large no-fishing areas, and four of the six fish groups that differed with management type were more abundant in larger no-fishing areas within a few years of establishment compared with small no-fishing and fished areas (data reported as statistical results). Fish assemblages were surveyed annually in 2002–2007, 2009 and 2012 at 16 sites, 9–16 m depth, in the Solitary Islands Marine Park. Four sampling sites were in each of four management areas (two no-take and two fished): small, no-fishing (<15 ha, established 1991), large, no fishing (>100 ha, established 2002), recreational fishing but no commercial fish trapping (>200 ha), and recreational fishing and commercial fish trapping (>200 ha per site). At each site, fish were recorded along six underwater visual transects (125 m2) and during three replicate five-minute timed-swim counts (250 m2).

    Study and other actions tested
  70. A study in 2008–2010 of an area of reef in a coastal marine park in the Indian Ocean, off western Australia (Speed et al. 2016) found that the time sharks spent inside a small protected area of the park where all fishing was prohibited for 20 years varied between three species and that immature sharks were more likely to remain in the protected area than adults, and thus receive more protection from fishing. Sicklefin lemon sharks Negaprion acutidens spent 98–99% of time in the no-fishing protected areas, blacktip reef sharks Carcharhinus melanopterus 0–99% and grey reef sharks Carcharhinus amblyrhynchos less than 1% of time. Immature sharks were located inside the no-fishing areas for 84–99% of time and adults 0–99%. In addition, immature sharks moved within smaller areas (0.6–8.5 km2) than adults (3.6–21.8 km2). Ningaloo Reef is the largest fringing reef in Australia (260 km long) and is protected by the multiple-use Ningaloo Marine Park established in 1987. Commercial fishing is prohibited and there are 18 no-fishing marine protected areas (884 km2). Sharks were caught and tagged in the marine park at beaches inside (by handlines) and outside (by longlines) a no-fishing area (11.35 km2) in February 2008 and November 2009. A total of 56 acoustic receivers deployed inside and outside the no-fishing area recorded tagged shark location every 30 minutes for up to two years. The movement data for 12 sharks consistently detected for six months or more were analysed.

    Study and other actions tested
  71. A site comparison study in 2008 of two submerged rocky cliff areas in the Tyrrhenian Sea, Italy (Appolloni et al. 2017) found higher overall fish species richness, higher fish abundance and biomass overall and for fisheries target species, and similar abundance and biomass of non-target fish inside a no-take zone of a marine protected area where all fishing was banned, compared to fished areas, although the effect varied with depth. Overall fish species richness, total abundance and total biomass were higher inside the no-take zone than fished areas at all depths (species richness: 14–18 vs 9–9, abundance: 235–357 fish vs 125–141 fish, biomass data reported as log-transformed). The abundance of targeted fish species was higher inside the no-take zone than fished areas at shallower depths (5 m: 136 vs 30, 10 m: 194 vs 25) but not at the deepest (20 m: 41 vs 23), and biomass was higher inside at all depths (data log-transformed). Abundance and biomass of non-target fish species were similar between areas (data reported as statistical results). The marine protected area at Punta Campanella (1,300 ha, year of designation not reported) extends from the coastline to 60 m depth and has two no-take areas. Underwater visual censuses were undertaken in June and October 2008 at one of the no-take zones where all fishing is banned (21 ha), and six partially protected sites where only some fishing (local fishers only and small vessels <10 gross tonnage) and other activities like diving are allowed. Fish were recorded along transects (25 m × 5 m × 5 m) at three depths (5 m, 10 m, 20 m). Three replicate transects were surveyed at each depth.

    Study and other actions tested
  72. A site comparison study in 2010–2011 at 12 coral reef sites in the Sulu Sea, Malaysia (Chung et al. 2017) found that prohibiting all fishing in a marine reserve resulted in a greater total fish density and biomass, and a higher biomass but similar density of coral trout Plectropomus spp., compared to fished areas outside, 11 years after implementation. Overall, areas closed to fishing had a higher reef fish density (closed: 624 fish/250 m2, fished: 373 fish/250 m2) and biomass (closed: 40 kg/250 m2, fished: 12 kg/250 m2) than fished areas. Average coral trout biomass was greater in closed (1.3 kg/250 m2) compared to fished areas (0.1 kg/250 m2), but density was similar (closed: 1.5 fish/250 m2; fished: 0.4 fish/250 m2). Sugud Islands Marine Conservation Area (467 km2) was established in December 2001 and prohibits fishing. Between April 2010 and November 2011, twelve patch reefs around Lankayan Island were surveyed: eight reefs in the reserve area closed to fishing and four open to fishing outside (0–3 km from the reserve border). Fish >3 cm length were recorded (count and species) by diver visual census along four randomly placed belt transects, 5 m wide by 50 m length, at each reef site (minimum 50–100 m apart). Fish biomass was estimated using length–weight relationships.

    Study and other actions tested
  73. A site comparison study in 2015 of 22 estuaries in the Tasman Sea, Australia (Gilby et al. 2017) found that prohibiting all fishing in estuarine reserves for between seven-12 years resulted in greater abundance of two of two non-harvested fish species, but lower abundance of four of four commercially harvested fish, compared to fished areas. Average abundance of species not harvested in the region (estuary perchlet Ambassis marianus and blue catfish Neoarius graeffei) was higher in unfished estuarine reserves than fished estuaries (unfished: 0.48–9.57 ind, fished: 0.12–6.33 ind), whereas average abundance of four fisheries-targeted species (yellowfin bream Acanthopagrus australis, grey mullet Mugil cephalus, common toadfish Marilyna pleurosticta, weeping toadfish Torquigener pleurogramma) was lower in the unfished reserves (unfished: 0.06–3.16 ind, fished: 0.39–8.99 ind). In addition, fish communities were different between unfished reserve and fished estuaries (data reported as a graphical analysis). The authors noted that differences in the environmental attributes between the unfished and fished estuaries contributed to the lower abundance in unfished estuaries for harvested fish. Data were collected between June and August 2015 in six no-take (no extractive activities, including fishing; one established 1993, the rest 2008) and 16 fished estuaries in the Moreton Bay Marine Park (created 1993). Fish were surveyed twice (for 1 h) over two days at ten sites (>250 m apart) in each estuary by baited remote underwater video. Counts were made of the maximum numbers of individual fish visible by species.

    Study and other actions tested
  74. A site comparison study in 2014 at two rocky sites in a bay in the North Sea off western Norway (Halvorsen et al. 2017) found that the commercial fishing mortality of corkwing wrasse Symphodus melops originally tagged at a small temporary marine protected area where fishing is prohibited was reduced compared to wrasse tagged at a fished site, but there was a similar selective removal of fish by size and sex regardless of site of origin. Overall fishing mortality of wrasse tagged inside the no-fishing site was lower than wrasse tagged in a fished site (not fished: 6–9%, fished: 31–41%). However, fishing mortality of nesting male wrasse (not fished: 12–15%, fished: 36–49%) was higher than for females (not fished: 3–5%, fished: 29–36%) at both sites. Average total length of nesting males was 119–141 mm and females 131–136 mm. In 2014, a total of 1,057 corkwing wrasses were tagged during (May–June) and after (July) the spawning period: 492 within a temporary no-fishing site (600 m of coastline, duration of protection was not reported) and 565 in a nearby site with no fishing restrictions. Fishing mortality of tagged wrasse was determined by recording the numbers captured and retained on all commercial potting fishing trips occurring within the bay over a three-month period.

    Study and other actions tested
  75. A site comparison study in 2007–2013 of 23 coral reef sites inside a marine reserve in the Caribbean Sea off Belize (Tewfik et al. 2017) found that the effects of prohibiting all fishing for 14–20 years on fish density, biomass and size varied with level in the food chain of five representative fish species/groups, compared to fished reserve zones. Data were presented graphically and as statistical results. Trends over time showed increases in average fish densities (fish/ha), biomasses (g/ha) or sizes (length to tail fork, mm) in the unfished zone compared to the fished zone: for large, and small, plant/algae-eating fish (Scaridae spp.), one invertebrate-eating fish (hogfish Lachnolaimus maximus) and two of three predatory (fish and/or invertebrates) fish (Nassau grouper Epinephelus striatus and black grouper Mycteroperca bonaci). Average density and biomass of the other predatory species (mutton snapper Lutjanus analis) showed no clear trends over time in the unfished zone, but size decreased. Black grouper density decreased, and biomass remained steady in both the unfished and fished zones, and small herbivores decreased in both unfished and fished areas over time. Diffferences between the unfished and fished zones were generally greater for the species at lower levels of the food chain (e.g. plant/algae eaters). Glover’s Reef Atoll was established as a Marine Reserve in 1993 and has several management zones including no-take (80 km2, all fishing banned), and general use (270 km2, fishing permitted, with regulations – see original paper for details). Fish in five no-take patch reefs and six fished reefs were monitored between 2007–2009, increased in 2010–2013 to include 12 additional fished reefs. Each reef was sampled once a year during April, May, or June. Fish number and estimated size over the entire area of each reef down to 3 m was recorded by snorkellers.

    Study and other actions tested
  76. A before-and-after study in 2007–2011 of reef and lagoon areas of an inhabited coral reef island in the Pacific Ocean, Tonga (Wenster et al. 2017) found that in the five years following the creation of a no-take fishing zone in a newly co-managed area that also excluded fishers from outside areas, the total fish catch rates in landed catches from the co-managed area did not increase, catch rates of half of the six individual species groups decreased and there was no decrease in overall fishing effort. No differences in total fish catch rates and catch rates of three of six fish groups (Acanthuridae - Naso spp., Holocentridae, Lethrinidae) were found since implementation, but catch rates of the remaining three (Acanthuridae - Acanthurus spp., Scaridae, Serranidae) decreased (data reported as statistical results). In addition, no difference in overall fishing effort was found (data reported as statistical results), but the authors reported that this was likely to be due to reduced travel to fishing grounds further away by resident fishers with the new exclusive rights. Co-management formally commenced on the island of ‘O’ua (one of 170 Tongan Islands) in 2007, covering a marine area of 4,606 ha, of which 203 ha is a no-take zone. Only residents on ‘Ou’a can fish the co-managed area. Fish catches were sampled (species and weight per trip) each year between 2007–2011 (total 184 records), collected opportunistically from landings by individual fishers (see original paper for fishing types). Catch data from spearfishing only was used for statistical analysis.

    Study and other actions tested
  77. A site comparison study in 2006–2013 of a large area of coral reef atolls in an island chain in the Pacific Ocean, USA (White et al. 2017) found that commercial fishing mortality of grey reef sharks Charcharhinus amblyrhnychos tagged at a large marine reserve where all fishing was banned for at least five years appeared to be low, and most of the sharks tracked by satellite remained inside the reserve, while some moved over large distances outside. The data were not statistically tested. Only 2% (five) of the 262 conventional tags deployed on sharks were recovered (the rest were either not caught or not reported), captured by small-scale fisheries at locations outside the reserve between 223–366 km away on average 587 days after tagging. Four of six sharks tracked by satellite remained inside the reserve boundaries for the entire monitoring period (1.3 years), and two were detected outside the reserve for 9% and 57% of time, travelling distances of up to 88 and 810 km respectively. Recovery of satellite-tagged sharks was not reported. Palmyra Atoll National Wildlife Refuge (54,126 km2) was established in 2001 (boundaries extended in 2009 and 2014) and prohibits all fishing and other extractive activities. From October 2006 to July 2009, a total of 262 grey reef sharks were caught in the reserve and marked with conventional tags on the dorsal fin. Recovered tags were actively sought (in 2007, 2009 and 2013) and encouraged from fishers at the three nearest inhabited and fished atolls several hundred kilometres away. During the same period, 11 fin-mounted satellite tags were deployed on adult sharks at the reserve, providing adequate data on the movements of six.

    Study and other actions tested
  78. A site comparison study in 2015–2016 of five areas of mixed seabed type (sand/seagrass/mangrove) in the Tasman Sea, eastern Australia (Henderson et al. 2018) found that over 15 months, giant shovelnose rays Glaucostegus typus spent more than half of the time inside marine reserves where all fishing was prohibited compared to fished areas outside, and it varied seasonally. Data were not statistically tested. Overall, rays were detected inside no-fishing areas compared to fished areas 58% of the days. In addition, ray detections inside the no-fishing reserves varied with season (winter: 53%, rest of year: 23%). Shovelnose rays were tracked from January 2015 to March 2016 in and around five marine reserves (fishing and extractive activities prohibited; the year established was not reported) located in Moreton Bay Marine Park. A total of 20 rays were surgically fitted with acoustic transmitters and released at two seagrass sites adjacent to reserves. The rays were tracked by 28 receivers (covering 180 km2) located in no-fishing and fished areas. Tracking data were analysed for 16 rays detected by receivers for longer than seven days, up to the removal of the receivers fifteen months later.

    Study and other actions tested
  79. A review of 10 studies of the effectiveness of different types of marine protected areas (study areas were not reported) (Sala & Giakoumi 2018) found that the total biomass of fish populations was highest in no-take marine protected areas relative to adjacent partially protected (some fishing permitted) marine protected areas and openly fished unprotected areas. The biomass of the whole fish assemblage was on average 670% greater within no-take protected areas than unprotected areas, and 343% greater than in partially protected areas. Fish biomass in partially protected areas was 183% greater than unprotected areas and was often similar. In addition, recovery of fish biomass over time was found in no-take areas after protection (nine–19 years), but not in partially protected or unprotected areas (data presented as log-ratios). The review surveyed peer-reviewed studies (total number not reported) documenting the biomass of whole fish assemblages of no-take marine reserves, partially protected marine protected areas, and open access areas all within the same vicinity. A meta-analysis of seven published and three unpublished studies (author and year reported only) comparing biomass data between all three areas was done.

    Study and other actions tested
Please cite as:

Taylor, N., Clarke, L.J., Alliji, K., Barrett, C., McIntyre, R., Smith, R.K., and Sutherland, W.J. (2021) Marine Fish Conservation: Global Evidence for the Effects of Selected Interventions. Synopses of Conservation Evidence Series. University of Cambridge, Cambridge, UK.

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