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Action: Designate a Marine Protected Area with a zonation system of activity restrictions Subtidal Benthic Invertebrate Conservation

Key messages

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  • Thirteen studies examined the effects of designating a marine protected area with a zonation system of activity restrictions on subtidal benthic invertebrate populations. Four studies were in the Caribbean Sea (Belize, Mexico), three in the Mediterranean Sea (Italy), one in the Central Pacific Ocean (Ecuador), three in the Bristol Channel and the Irish Sea (UK), one in the Indian Ocean (Australia), and one in the North Atlantic Ocean (Portugal).

 

COMMUNITY RESPONSE (2 STUDIES)

  • Overall community composition (1 study): One site comparison study in the Mediterranean Sea found that inside a marine protected area with a zonation system, the combined invertebrate and algae species community composition was different at a site prohibiting all fishing compared to sites where some fishing occurs, after six years.
  • Overall species richness/diversity (1 study): One site comparison study in the North Atlantic Ocean found that inside a marine protected area with a zonation system, sites prohibiting nearly all fishing had similar invertebrate species richness to sites where fishing was mostly allowed, after two years.

POPULATION RESPONSE (13 STUDIES)

  • Overall abundance (1 study): One site comparison study in the North Atlantic Ocean found that inside a marine protected area with a zonation system, abundances of specific invertebrate groups varied between sites prohibiting nearly all fishing and sites where fishing was mostly allowed, after two years.
  • Crustacean abundance (7 studies): Three of seven site comparison studies (two replicated) in the Caribbean Sea, the Central Pacific Ocean, and in the Bristol Channel and the Irish Sea found that inside a marine protected area with a zonation system, abundance and/or biomass of spiny lobsters increased in a zone closed to all/commercial fishing and were greater than in a zone where fewer fishing restrictions occurred, after four to 20 years depending on the study. One found that sites closed to all fishing had higher abundances of spiny lobsters and slipper lobsters after eight to ten years compared to fished sites. Two found that sites closed to all fishing for six to seven years had more European lobsters than sites where potting was allowed. And one found that abundances of European lobsters, velvet crabs, brown crabs and spider crabs, after one to four years, varied with the levels of protection.
  • Crustacean condition (4 studies): Three of five site comparison studies (one replicated) in the Bristol Channel and the Irish Sea, and in the Caribbean Sea found that, inside a marine protected area with a zonation system, sites prohibiting all fishing for seven years or commercial fishing (duration unspecified) had bigger lobsters compared to fished areas. One found that the sizes of lobsters, velvet crabs, brown crabs and spider crabs varied with the levels of protection, and one study found that the size of spiny lobsters decreased similarly in an area prohibiting all fishing and in an area with fewer restrictions 14 to 20 years after designation of the protected area. Two studies undertaken in the same area found conflicting effects of prohibiting all fishing for six to seven years on disease and injury of lobsters.
  • Echinoderm abundance (2 studies): One of two site comparison studies in the Mediterranean Sea found that inside a marine protected area with a zonation system, at a site prohibiting all fishing for 17 to 18 years, abundances of two species of sea urchins were higher than at sites allowing the recreational fishing of purple sea urchins. The other one found similar abundance of purple sea urchins inside fully protected sites, sites where some restricted urchin harvest occurs, and unprotected fished sites outside the protected area after five years.
  • Echinoderm condition (2 studies): Two site comparison studies in the Mediterranean Sea found that inside a marine protected area with a zonation system, sites prohibiting all fishing had bigger sea urchins compared to sites where some restricted urchin harvest occurs and compared to unprotected fished sites outside the protected area, after either four years or 17 to 18 years.
  • Mollusc abundance (3 studies): One replicated, randomized, controlled study in the Indian Ocean found that inside a marine protected area with a zonation system, abundance of blacklip abalone was higher in sites that had been prohibiting all fishing for five years compared to those prohibiting commercial fishing only. Two site comparison studies in the Caribbean Sea found that inside marine protected areas with a zonation system, abundances of adult queen conch increased over time in a zone closed to all fishing and were greater than in zones with fewer restrictions, but abundances of juvenile conch did not differ or vary differently between zones, after either five to eight years or 14 to 20 years.
  • Mollusc condition (1 study): One site comparison study in the Caribbean Sea found that inside a marine protected area with a zonation system, the size of queen conch decreased similarly in the area prohibiting all fishing and in the area with fewer restrictions, after 14 to 20 years.
  • Sponge abundance (1 study): One site comparison study in the Mediterranean Sea found that inside a marine protected area with a zonation system, the cover of sponges Cliona was higher at a site prohibiting all fishing for six years compared to sites where some fishing occurred.

BEHAVIOUR (1 STUDY)

  • Crustacean behaviour (1 study): One site comparison study in the Caribbean Sea found that, inside a marine protected area with a zonation system (year of designation unspecified), 80% of the lobster population occurring in the unfished area remained in the protected unfished area, and thus remained protected.

Supporting evidence from individual studies

1 

A site comparison study from 1997–2001 of two coral reef, seagrass and sandy seabed areas in Glover’s Reef marine reserve, western Caribbean Sea, off the coast of Belize (Acosta 2002) found that between five and eight years after designating a marine protected area with a zonation system, abundances of adult spiny lobster Panulirus argus and adult queen conch Strombus gigas increased in a zone closed to all fishing and were greater than in a zone where commercial fishing occurred. Abundance of adult lobsters (>45 mm carapace length) increased in the closed zone (after four years in 1997: 21; after eight years in 2001: 84 lobsters/ha) and was greater than in the fished zone where abundance did not change (1997: 13; 2001: 26). Abundance of adult conch (>110 mm shell length) increased in the closed zone (1997: 244, 2001: 921 conch/ha), and was greater than in the fished zone where abundance did not change (1997: 296, 2001: 188). Abundance of juvenile lobsters and conch did not vary over time or differ between zones. The reserve was established in 1993, with a general use zone open to commercial fishing and a zone prohibiting all fishing. In each zone inside the reserve, divers counted and measured lobsters (in eight 0.5 ha coral reef patches/zone) and conch (in twenty-four 200 m2 transects on sand/zone) at quarterly intervals in 1997–2001.

2 

A site comparison study from 1996–2001 of two coral reef, seagrass, and sandy seabed areas at Glover’s Reef marine reserve, western Caribbean Sea, off the coast of Belize (Acosta & Robertson 2003) found that between four and eight years after designating a marine protected area with a zonation system, abundance and biomass of Caribbean spiny lobster Panulirus argus increased in a zone closed to all commercial fishing and were also greater than in a zone where artisanal commercial fishing occurred. Annual average abundance of lobsters increased in the closed zone (after three years in 1996: 63, after eight years in 2001: 144 lobsters/ha), and was greater than in the fished zone where abundance did not change (1996: 42; 2001: 61 lobsters/ha). Biomass of adult lobsters (>76 mm carapace length) increased in the closed zone (1996: 10; 2001: 155 kg/ha) and was higher than in the fished zone where biomass did not change (1996: 7; 2001: 3.5 kg/ha). All data were extracted from the graphs presented in the study. The reserve was established in 1993, with a general use zone open to artisanal commercial fishing and a zone prohibiting all commercial fishing. In each zone inside the reserve, divers counted and measured lobsters in eight 0.007–0.5 ha coral reef patches/zone at quarterly intervals in 1996–2001.

3 

A site comparison study in 2002–2003 of three rocky sites within a marine protected area with a zonation system in northeast Sardinia, Mediterranean Sea, Italy (Ceccherelli et al. 2006) found that a fully protected site prohibiting all fishing had a different invertebrate and algae (species combined) community composition to partially protected sites where some fishing occurred, after six years of enforcement. Community data were reported as graphical analyses. In addition, the cover of sponges Cliona spp. was higher in the fully protected site (6–12 % cover) compared to the partially protected sites (1–5 % cover). In September 2002 and 2003, algae and invertebrates were surveyed once at three sites (5 m depth) inside a protected area: a no-take zone (one site) and a partially protected zone where some regulated fishing takes places (two sites). Restrictions and limitations had been enforced since 1997. Divers photographed 10 quadrats (16 × 23 cm)/site. Percentage cover of sessile invertebrates and algae were estimated from photographs.

4 

A site comparison study in 2003–2004 of three rocky seabed sites within a marine protected area with a zonation system, off Ustica Island, Mediterranean Sea, Italy (Gianguzza et al. 2006) found that overall at a fully protected site that had been prohibiting all fishing for 17–18 years, abundances and sizes of two species of sea urchins were higher than at partially protected sites where recreational fishing of purple sea urchins Paracentrotus lividus occurred, but effects varied seasonally. Abundances of purple and black sea urchins Arbacia lixula were higher in the fully protected than the partially protected sites in summer (purple: 2.9 vs 0.7–1.3/m2, black: 3.1 vs 1.7–1.9/m2) and autumn (purple: 4.1 vs 1.6–2.1/m2, black: 2.3 vs 0.7–1/m2), but not spring (purple: 2.3 vs 2.3–2.9/m2, black: 1.5 vs 2.0/m2). Purple sea urchins were larger in the fully protected than the partially protected sites in spring (fully: 45 vs partially: 31–34 mm), summer (43 vs 35–37 mm) and autumn (44 vs 32–35 mm). Black sea urchins were smaller in the fully protected than the partially protected sites in autumn (31 vs 35–37 mm), but similarly sized across sites in spring and summer (37–42 vs 39–45 mm). Ustica Island marine protected area was established in 1986 with a no-take zone and a partially protected zone where some recreational activities take place. In 1994, recreational fishing for purple sea urchin inside the partially protected zone was reopened following undesirable increases in their abundance leading to barren areas. At one site in the no-take zone and two in the partially protected (4–8 m depth), divers identified and counted all urchins along three 10 m2 transects, twice in autumn 2003, in spring 2004, and in summer 2004. The diameter (not including spines) of urchins inside 1 m2 quadrats was measured.

5 

A site comparison study in 2006 of seven sites in a seagrass and rocky seabed area in the Mediterranean Sea, Sardinia, Italy (Ceccherelli et al. 2009) found that the effect of designating a marine protected area with a zonation system on purple sea urchin Paracentrotus lividus abundance and size varied with the level of restriction in place. Within the protected area after four years, fully protected no-take sites had similar abundances of urchin (2–5 individuals) compared to partially protected sites where some restricted urchin harvest occurred (1–12 individuals), and to unprotected fished sites outside the protected area (2–12 individuals). However, urchins were larger in no-take sites (57–62 mm), compared to partially protected (32–61 mm) and unprotected fished sites (24–50 mm). Capo Caccia–Isola Piana marine protected area was established in 2002 with varying levels of protection including a no-take zone and a partially protected zone where urchin harvest was formerly prohibited but reopened with restrictions in 2006 (see paper for details). Sampling took place in April–May 2006 after the harvest season at seven sites (200 m2 each) in 6–10 m water depth: one within the no-take zone, three within the partially protected zone, and three outside the marine protected area. At each site, urchins were counted inside 20 quadrats (1 × 1 m), and 20 urchins were measured (diameter without spines).

6 

A replicated, site comparison study in 2000–2002 of 20 rocky seabed sites inside the Galapagos Marine Reserve, eastern Pacific Ocean, Ecuador (Sonnenholzner et al. 2009) found that protected sites that had been closed to all fishing for eight to ten years had higher abundances of spiny lobsters Panulirus penicillatus and slipper lobsters Scyllarides astori, compared to fished sites inside the reserve. Encounter rates (indicative of abundance) of spiny lobster and slipper lobster were higher in the closed areas (spiny: 0.4; slipper: 0.2 lobsters/hr) than the fished areas (spiny: 0.1; slipper: 0.1 lobsters/hr). Pencil urchin Eucidaris galapagensis abundance was lower in closed areas (2.2 urchins/m2) than fished areas (4.5 urchins/m2). Fishing exclusion zones within the reserve were created in 1992 and formally established in 2000, but uneven compliance with the fishing regulations is reported. In April 2000–August 2002, divers surveyed lobsters and sea urchins at ten sites within exclusion zones and ten sites outside (but inside the reserve). Lobsters were counted along four 20-min dive transects/site.

7 

A replicated, site comparison study in summer 2004–2007 of ten rocky and sandy sites, across two zones inside a marine protected area and two areas outside, in the Bristol Channel and the Irish Sea, UK (Hoskin et al. 2011) found that abundances and sizes of European lobster Homarus gammarus and three crab species   varied with the levels of protection. Abundance of large lobsters (≥90 mm) increased by 127% inside the no-take zone between 2004 and 2007 (one to four years after designation of the no-take zone; from 3 to 7 lobsters/line) and was five times higher than in aa partly fished zone (potting only) inside the protected area and fully fished areas outside where abundance had not changed (1–2 lobsters/line). Abundance of small lobsters (<90 mm) increased by 97% in the no-take zone (from 3 to 7 lobsters/line) and by 140% in the potting-only zones (argued by the authors to be due to spill-over effects; from 2 to 4–7 lobsters/line), where they appeared greater than in the fully fished areas where abundance remained constant (2–4 lobsters/line). The size of large lobsters (≥90 mm) increased by 5% inside the no-take zone between 2004 (98 mm) and 2007 (103 mm) and became 9% larger than in the potting-only zone and fished areas where lobster size decreased by 2–3% (from 98 to 95 mm). The size of small lobsters did not change over time and was similar across all areas. Abundance of velvet crabs Necora puber decreased by 65% inside the no-take zone over time (from 2 to 1 crab/line; argued by the authors to be due to increased predation by lobsters) and decreased even more in the potting-only zones (from 5–6 to 1 crab/line), and appeared lower than in the fully fished areas where it increased (from 0–6 to 1–7 crabs/line). The average size of velvet crabs did not change over time and was similar across all areas. Abundance of brown crabs Cancer pagurus did not change over time inside the no-take zone (0.3 crab/line), nor in the potting-only zone and fished areas (from 0.3–2 crabs/line). The average size of brown crabs increased by 25% inside the no-take zone between 2004 (115 mm) and 2007 (144 mm) and became greater than in the potting-only zones (123–128 mm) but not in fully fished areas (116–130 mm). Abundance of spider crabs Maja squinado was similar in 2004 and 2007 for all areas but varied spatially (with the no-take zone having lower abundance). The average size of spider crabs did not change over time and was similar across all areas. Lundy Island marine protected area was designated as a voluntary reserve in 1971 (statutory since 1986). In 2003, it included a 4 km2 no-take zone (no fishing or harvesting allowed), the rest being a refuge zone only allowing crab and lobster potting. In 2004–2007, lobsters and crabs were surveyed inside the no-take zone, at two locations in the refuge zone, and two distant fished locations (20–100 km away) (2 sites/location). Four lines of standard commercial baited shellfish pots were deployed (10 pots/line) at each site for 24 h. Upon retrieval, lobsters and crabs were counted and measured (carapace length). The pots were redeployed for five consecutive days each year.

8 

A replicated, randomized, controlled study in 2008–2012 of nine rocky reef sites inside a marine park in Shark Bay, Indian Ocean, southeastern New South Wales, Australia (Wootton et al. 2012) found that five years after designating a marine park with various levels of fishing restrictions, the abundance of blacklip abalone Haliotis rubra was higher in sites with full fishing prohibition compared to those with partial prohibition and compared to sites outside the park (all fishing allowed). There were more abalone in sites with full fishing prohibition (4.3 individuals/transect) compared to sites with partial prohibition (0.9) and sites outside the park (1.9). In 2007, a marine park was established which included zones where all fishing was prohibited, and zones with partial prohibition (commercial fishing prohibited but recreational fishing and harvesting allowed). Twice annually between 2008 and 2012, samples were collected at nine randomly selected sites: three within each prohibition level inside the park, three outside the park where all fishing is allowed. Three 30 m transects/site were randomly deployed at 1–3 m depth, and abundance of blacklip abalone estimated from one 1 m strip/transect.

9 

A site comparison study in 2010 of six sites in two zones inside a marine protected area in the Bristol Channel, UK (Coleman et al. 2013 – similar set-up as Davies et al. 2014) found that sites in the no-take zone (where all fishing had been prohibited for six years) had more and bigger European lobsters Homarus gammarus than sites outside in the refuge zone where potting was allowed. Lobsters were caught in higher abundance inside the no-take zone (514) than outside (152) and grew bigger inside (99 mm) than outside (86 mm). In addition, more lobsters were above the minimum landing size (90 mm) inside the no-take zone (75% of lobsters) than in the refuge zone (36% of lobsters). A higher proportion of egg-bearing females were found in the no-take zone (31%) compared to the refuge zone (7%). Overall, similar proportions of injured lobsters were found inside the no-take zone (33%) and inside the refuge zone (26%). The percentage of diseased lobsters was higher inside the no-take zone (27%) compared to the refuge zone (18%). Lundy Island marine protected area was designated as a voluntary reserve in 1971 (statutory since 1986). In 2003, it included a 4 km2 no-take zone (no fishing or harvesting allowed), the rest being a refuge zone only allowing crab and lobster potting (all other fishing is prohibited). In 2010, six sites inside the protected area were surveyed: two within the no-take zone and four in the refuge zone. At each site, one line of 35 baited pots was deployed for 24–48 h, and all lobsters caught were measured (carapace length), sexed, assessed for injuries and diseases, and released back into the water. This process was repeated continuously over four days in May and again in June.

10 

A site comparison study in 2011 of six sites in two zones inside a marine protected area in the Bristol Channel, UK (Davies et al. 2014 – similar set-up as Coleman et al. 2013) found that sites in the no-take zone (where all fishing had been prohibited for seven years) had more and bigger European lobsters Homarus gammarus than sites outside in the zone where potting was allowed. Lobsters were more abundant inside the no-take zone (40/line of 35 pots) than outside (20), and grew bigger inside (93 mm) than outside (85 mm). In addition, more lobsters were above the minimum landing size (90 mm) inside the no-take zone (61% of lobsters) than outside (32% of lobsters). Because more and bigger lobsters occurred inside the no-take zone, more were found injured (inside: 41%; outside: 19%; assumed to be likely due to increases in fighting behaviour). The percentage of diseased lobsters was similar inside (28%) and outside (17%) the no-take zone. Lundy Island marine protected area was designated as a voluntary reserve in 1971 (statutory since 1986). In 2003, it included a 4 km2 no-take zone (no fishing or harvesting allowed), the rest being a refuge zone only allowing crab and lobster potting (all other fishing is prohibited). In August 2011, six sites inside the protected area were surveyed: two within the no-take zone and four outside in the refuge zone. At each site, one line of 35 baited pots was deployed for 24 h, and all lobsters caught were measured (carapace length) and assessed for injuries and diseases.

11 

A site comparison study in 2011–2012 of two areas within a marine protected area in the Caribbean Sea, Mexico (Ley‐Cooper et al. 2014) found that Caribbean spiny lobsters Panulirus argus grew larger in an area where commercial fishing was banned compared to a fished area, and that the majority of the lobster population in the unfished area remained protected. Lobster sizes were greater in the unfished area (94 mm) compared to the fished area (73 mm). In the unfished area, this corresponded to 99% of lobsters being bigger than the minimum legal catch size (74.5 mm), while in the fished area it corresponded to only 25%. In addition, an estimated 20% of the lobster population occurring in the unfished area moved to the fished area, thus 80% remained protected. The study was carried out in a Biosphere Reserve (year of designation unspecified) which restricted commercial fishing to shallow depths (<20 m) and banned it where depths exceed 20 m (see paper for details). In August–September 2011, lobsters were hand-caught from the unfished area, tagged, sized (carapace length) and released in the unfished area (379 in total). During the 2011/2012 fishing season in the fished area, all lobsters caught by fishermen were sized, and tagged lobsters recorded. A tag-recapture model based on the number of recaptured tagged lobsters (20 in total) was used to estimate the percentage of the lobster population moving from the protected to the fished area.

12 

A site comparison study in 2013 of four rocky seabed sites inside a marine park with a zonation system in the North Atlantic Ocean, southwest Portugal (Gil Fernandez et al. 2016) found that sites prohibiting nearly all fishing had similar invertebrate species richness to sites where fishing was mostly allowed, two years after implementation. Sites prohibiting nearly all fishing had six species and sites where fishing was mostly allowed had seven species. In addition, abundances of specific groups appeared to vary between sites prohibiting nearly all fishing and sites mostly allowing fishing (sea urchins: 7 vs 31; brittle stars: 4 vs 63; starfish: 0–8 vs 1–39; sea cucumbers: 12 vs 31; octopus: 1 vs 5; data not statistically tested; unit unspecified). Fishery restrictions inside the park were implemented in 2011. In February–May 2013, four partially protected sites were sampled (0–15 m depth): two where nearly all professional and recreational fishing were prohibited (only barnacle extraction permitted), and two where fishing was mostly allowed (bottom trawling and recreational fishing not allowed on Wednesdays). Divers identified and counted all macro-invertebrates (size unspecified) along four 10 × 2 m transects/site.

13 

A site comparison study in 2007–2013 of 11–23 coral reef sites inside Glover’s Reef Marine Reserve, Caribbean Sea, Belize (Tewfik et al. 2017) found that the effects of a protected no-take area on the abundances and sizes of queen conch Lobatus gigas and Caribbean spiny lobster Panulirus argus, compared to the protected general-use zone with only some restrictions, varied with the size of individuals., Inside the marine reserve, 14 to 20 years after its designation, abundance of mature conch (>5 mm lip thickness) increased over time in the no-take sites (from 4/ha in 2007 to 17/ha in 2013), and was greater than in the general-use sites where the change (from 1 to 2/ha) was not significant. Immature conch (<5 mm) abundance increased similarly in no-take (from 4 to 53/ha) and general-use sites (from <1 to 33/ha). The lip thickness of mature conch decreased similarly over time at all sites (from 11 to 9 mm). The shell length of immature conch decreased similarly over time in no-take sites (from 221 to 182 mm) and general-use sites (from 234 to 186 mm). Abundance of legal-size (>76 mm carapace length) and sub-legal (<76 mm) lobsters increased over time in the no-take sites (legal-size: from 6 to 16/ha; sub-legal: from 1 to 3/ha) but did not change in the general-use sites (legal-size: non-significant change from 6 to 5/ha; sub-legal: remained at 6/ha). The size of all lobsters decreased over time in both no-take sites (legal-size: from 120 to 110 mm; sub-legal: from 59 to 52 mm) and general-use sites (legal-size: from 110 to 100 mm; sub-legal: from 59 to 52 mm). Glover’s Reef Atoll was designated as a Marine Reserve in 1993 and included a no-take area (79.6 km2) and a general-use area with fishery restrictions (including: ban on the use of SCUBA to collect any seafood, closed seasons, and size limits for queen conch and spiny lobster). Once a year in April–June 2007–2013, conch and lobsters were surveyed at 1.6 m average depth inside the no-take area (6–18 sites/year) and inside the general-use zone (5 sites/year). At each site (0.04–1.43 ha), snorkelers counted and measured all conch (shell length; lip thickness) and lobster (carapace length).

Referenced papers

Please cite as:

Lemasson, A.J., Pettit, L.R., Smith, R.K., and Sutherland, W.J. (2019) Subtidal Benthic Invertebrate Conservation: Global Evidence for the Effects of Interventions. Synopses of Conservation Evidence Series. University of Cambridge, Cambridge, UK.