The potential value of indicator groups in the selection of marine reserves

Efforts to systematically select networks of marine reserves to conserve biodiversity may be constrained by limited information on the distribution of biodiversity. A potential solution is the use of surrogates, such as indicator groups, that effectively represent other elements of biodiversity. The potential of macroalgae and molluscs to act as indicator groups for all species in the selection of intertidal marine reserves was tested for a sample of rocky shore locations in south-east Australia. Fifteen locations were surveyed between November 1999 and January 2000, and reserve selection for each of the potential indicator groups and for all species was undertaken using richness-based and rarity-based algorithms. The sets of locations chosen by both algorithms to conserve each of the indicator groups included around 95% of all species. There was 54% spatial correspondence of the sets of locations chosen for each of the indicator groups. Molluscs appeared to be a reliable indicator for areas of reservation ranging from 6.7 to 73.3% of the locations surveyed, including a similar percentage of all species as the locations selected for all species. Unlike macroalgae, locations chosen for molluscs included significantly more species than a set of locations chosen at random, and more species than the set of locations chosen for randomly selected subsets of species of the same richness as molluscs. There is significant scope for further research into the value of indicator groups in reserve selection in marine systems.     

Do partial marine reserves protect reef fish assemblages?

Fish assemblages in the Mimiwhangata Marine Park, an area closed to commercial fishing but open to most forms of recreational fishing, were compared with adjacent fished areas. Two survey methodologies were used; baited underwater video and underwater visual census. Snapper (Pagrus auratus), the most heavily targeted fish species in the region, showed no difference in abundance or size between the Marine Park and adjacent control areas. When compared to the fully no-take Poor Knights Island Marine Reserve and two other reference areas open to all kinds of fishing (Cape Brett and the Mokohinau Islands), the abundance and size of snapper at the Marine Park were most similar to fished reference areas. In fact, the Marine Park had the lowest mean numbers and sizes of snapper of all areas, no-take or open to fishing. Baited underwater video found that pigfish (Bodianus unimaculatus), leatherjackets (Parika scaber) and trevally (Pseudocaranx dentex) were significantly more common in the Marine Park, than in the adjacent control areas. However, none of these species are heavily targeted by fishers. Underwater visual census found similar results with five species significantly more abundant in the Marine Park and five species more abundant outside the Marine Park. The lack of any recovery by snapper within the Marine Park, despite the exclusion of commercial fishers and restrictions on recreational fishing, indicates that partial closures are ineffective as conservation tools. The data suggest fishing pressure within the Marine Park is at least as high as at other ‘fished’ sites.     

Effects of fishing on sex-changing Caribbean parrotfishes

We studied parrotfish (Scaridae) assemblages on coral reefs in relation to fishing pressure around six Caribbean islands. Fishing intensity ranged from virtually none in Bonaire, and increased through Saba, Puerto Rico, St Lucia and Dominica to extremely high levels in Jamaica. In St Lucia we also compared parrotfish assemblages between fishing grounds and fully protected marine reserves, from 1995, 6 months prior to establishment, to 2001. Within each country we performed replicate counts of the number and size of all parrotfish species within, or passing through our counting area. From these data we calculated biomass for seven species. Biomass of the two largest species, Sparisoma viride and Scarus vetula, was greatest in islands with low fishing pressure (P<0.001). By contrast, smaller species constituted an increasing proportion of the total parrotfish assemblage as fishing pressure increased (P<0.001 in all cases). Parrotfish are protogynous hermaphrodites with two distinct colour phases. The initial phase is predominantly female, and the terminal phase exclusive to sexually mature males. The average size of all species except Sc. vetula tended to decrease with increasing fishing pressure. Furthermore, percentages of fish that were terminal phase males showed order of magnitude declines with increasing fishing pressure for Sp. viride and Sc. vetula. Terminal males of these species were absent from counts in Jamaica and virtually absent from Dominica suggesting that persistence of these populations may depend on recruitment from distant sources. Following reserve implementation in St Lucia, all species, except uncommon Sp. chrysopterum, increased in mean biomass (P<0.001 in all cases). In 6 years the total biomass for all species combined increased to become nearly four times as high in reserves and almost twice as high in fishing grounds [P<0.001 (year effect); P<0.001 (protection effect); P<0.001 (year×protection)], and mean size of five species increased significantly in both reserves and fishing grounds.     

The influence of life history attributes and fishing pressure on the efficacy of marine reserves

Two key questions regarding “no-take” marine reserves are: (1) how effective are reserves likely to be, and (2) how does effectiveness vary with life history attributes and the relative size of reserves. To investigate these questions, we use a simple Ricker model that includes fishing, larval dispersal, and larval loss while in a planktonic pool, and that tracks protected and unprotected populations. We applied two different measures of reserve effectiveness to our simulation results. One metric was intended to reflect goals oriented towards conservation and the second was intended to reflect fishery enhancement goals. Both metrics compare the situation before reserves are established to after the reserve has been in place and a new equilibrium was reached. Yield effectiveness is defined as the total equilibrium annual harvest after reserves are established divided by the total annual harvest before reserves are established. Conservation effectiveness is defined as the average adult density inside the reserve divided by the average density in the same area prior to reserve establishment. A substantial fraction of the 5120 simulated parameter combinations representing different harvest rates and life history attributes went extinct in the absence of a reserve, and these scenarios leading to extinction could be predicted accurately (85% aptly classified) simply on the basis of exploitation rate and population growth rate. Of the cases that did not go extinct, we compared the performance of reserves as measured by each effectiveness metric. Few of the cases (less than 8%) produced effective reserves as measured in terms of increased harvest; whereas over half of the cases resulted in effective reserves as measured by conservation effectiveness. Moreover, the two measures of reserve effectiveness were only weakly correlated. Simple linear regression or polynomial regression could explain at most 23% of the variation in reserve effectiveness as measured by either metric. As expected, the size of the reserve area had a marked and typically negative effect on total annual yield, which suggests that while marine protected areas may do a good job of conserving protected populations, there will generally be pressure from the fishing community to keep them small because of their tendency to reduce total catch.     

Persistence of depleted abalones in marine reserves of central California

We evaluated the persistence and possible recovery of two depleted marine mollusks, the red (Haliotis rufescens) and black abalone (H. cracherodii), in central California, USA. Monitoring over 32-years did not reveal increasing or decreasing trends in red abalone abundances and sizes over the past three decades, in the absence of harvesting. Comparisons between marine reserves protected for at least 25 years and sites with open public access showed significant difference in size structure for black abalone, with individuals greater than 8 cm in shell length comprising 14–37% of animals in reserves and 2–11% at open-access sites, and a trend for greater abundances of red abalone within reserves. Despite no increasing trends, protection in one of the no-take reserves, the Hopkins marine life refuge (HMLR), has led to persistence of red abalone populations over multiple generations, at average densities of 0.2 individuals/m². At other locations, both within the HMLR and elsewhere, red abalone densities are lower than at the location where long-term studies were conducted (av. 0.03 animals/m²), and an order of magnitude lower than for black abalone (av. 0.4 animals/m²). These results suggest that continued fishery closure and protection in no-take reserves are effective tools for allowing persistence of abalone populations, though there are no signs of recovery to levels comparable to those preceding fisheries collapse. Such failure to recover is most likely associated with high natural mortality and possibly continued illegal take, but not with processes underlying low abalone population levels elsewhere, including food or habitat limitation, recruitment failure, or disease. Linking current structure and trends to specific processes is a crucial first step towards devising focused strategies for conserving and re-building depleted marine populations.     

Conservation of coral reef biodiversity: a comparison of reserve selection procedures for corals and fishes

A range of different biodiversity-based selection methods for nature reserves has been tested for terrestrial environments, including those based on diversity hotspots, endemicity hotspots and complementarity. In this study, we investigate the utility of these approaches for a coral reef embayment. We compare coral and fish species richness in a random accumulation of reserve sites with (a) hotspots analysis, (b) stratified selection of hotspots, and (c) complementarity. Cumulative species-site curves indicated that complementarity maximized the rate of accumulation of species of both corals and fishes in reserves, while the hotspot approach performed moderately well. An equivalent number of reserve sites supported a greater proportion of the coral biodiversity when compared to fishes, reflecting the broader distribution of corals. Our results indicate that when choosing an indicator group as a proxy for representing overall diversity in a reserve network, the group with the greatest heterogeneity will provide the best results. Our findings also show that although a modest number of protected sites (20%) will incorporate much of the local diversity (>75%), species-specific approaches must be incorporated to target rare species.     

Connectivity of reef fish between mangroves and coral reefs: Algorithms for the design of marine reserves at seascape scales

Many species of coral reef fish undertake ontogenetic migrations between seagrass beds, mangroves, and coral reefs. A recent study from the Caribbean found that the availability of mangrove nursery habitat had a striking impact on the community structure and biomass of reef fish in their adult, coral reef habitat. The biomass of several species more than doubled when the reefs were connected to rich mangrove resources (defined as having at least 70 km of fringing Rhizophora mangle within a region of 200 km²). Here, the results of this large-scale empirical study are translated into a series of algorithms for use in natural resource management planning. Four algorithms are described that identify (i) the relative importance of mangrove nursery sites, (ii) the connectivity of individual reefs to mangrove nurseries, (iii) areas of nursery habitat that have an unusually large importance to specific reefs, and (iv) priority sites for mangrove reforestation projects. The algorithms generate a connectivity matrix among mangroves and coral reefs that facilitates the identification of connected corridors of habitats within a dynamic planning environment (e.g., reserve selection algorithms).     

Links between sex change and fish densities in marine protected areas

Sex change is widespread among marine fishes, including many species that are fished heavily, and is thought to be of conservation concern under some circumstances. As such, an important question in conservation is whether the implementation of marine protected areas (MPAs), which is a commonly used marine conservation tool, works as effectively for sex-changers as for non-sex-changers. To address this issue, we used meta-analyses of the ratio of fish abundances inside vs. outside MPAs to determine whether sex change affects the extent to which fish densities respond to protection. When all data were considered, there were similar responses to protection irrespective of reproductive mode. However, when analyses were restricted to older reserves (at least 10 years’ protection), female-first sex-changers consistently benefited from protection. Non-sex-changers and male-first sex-changers showed more variable responses to protection and, as a result, there were no significant differences between fish with different reproductive modes in their overall response to protection. The same results were observed when the effects of fisheries status (targeted vs. not targeted) were controlled. Our results support the use of MPAs as important components of conservation and demonstrate that old reserves are most consistently beneficial to female-first sex-changing species. Finally, our results highlight the fact that some effects of protection are only detectable after several generations.     

The role of social aggregations and protected areas in killer whale conservation: The mixed blessing of critical habitat

Protected areas are often proposed as tools for conserving endangered populations of marine megafauna. Our study area includes a voluntary no-entry reserve embedded within wider critical habitat for Threatened ‘northern resident’ killer whales under Canada’s Species at Risk Act. Our study quantified the reserve’s importance to whales by assessing habitat preference in a behavioural context, and population-level implications of that preference given threats from human activities, such as oil spills resulting from shipping traffic. We recorded summertime activities of whales from 1995 to 2002. Whales were observed on 397 of 530 (74.9%) days. Whales showed strong preference for the reserve over adjacent waters, and used it preferentially for feeding and beach-rubbing. While the area comprises ∼0.001% of the whales’ range, an overall average of 6.5% of the population was present each day. Frequently, >50% of this small population was aggregated in the restricted and heavily trafficked waterway of Johnstone Strait. Using the Potential Biological Removal equation, we calculated potential annual mortality limits (ML) of 2.2 animals. Mean group size in the area exceeded ML on 55.8% of days overall, and 98.8% of days when conditioning on whale presence. The whales’ high reliance on a trivial fraction of their range means that opportunities are routine for one stochastic, catastrophic event to cause population decline. On 20 August 2007, a barge loaded with ∼10,000 L of diesel sank in the area, exposing 25% of the population. This underscores the importance of identifying critical habitat for threatened populations, and ensuring meaningful protection.     

Evidence for a depth refuge effect in artisanal coral reef fisheries

Natural refuges can play a fundamental role in protecting species from overexploitation but have not been adequately quantified in the marine environment. We quantified the effect of a depth refuge on all fish species in an artisanal coral reef fishery in Zanzibar, Tanzania by comparing changes in fish species richness and relative abundance with depth at five fished and three unfished reefs across the region. Commercial species richness was depleted by 15.1% at shallow depths in fished reefs, but there was no difference between the reefs deeper than seven metres. Non-commercial species were not affected by fishing or depth. Evidence for similar patterns in fish communities in other countries and depth-limitations to artisanal fishing methods imply this effect is widespread. The depth refuge effect could be sustaining coral reef fisheries and should be taken in to account before implementing policies or subsidies that encourage or allow fisheries to exploit deeper waters.     

Testing the utility of abiotic surrogates for marine habitat mapping at scales relevant to management

Habitat mapping at the scale at which marine protected areas are designed and managed is essential for assessment of, and design for, representation. Most habitat mapping studies rely solely or in part on abiotic surrogates for patterns of biodiversity. The utility of abiotic variables in predicting biological distributions at the local scale (10 s of km) was tested in a remote video survey of macrobenthos in Moreton Bay, Australia. Habitat classifications of the same set of 41 sites based on 6 abiotic variables and abundances of 89 taxa and bioturbation indicators were compared using correlation, regression and ordination analyses. The concepts of false homogeneity (abiotically similar but biologically distinct) and false heterogeneity (abiotically distinct but biologically similar) were defined to describe types of errors associated with using abiotic surrogates to construct habitat maps, and quantified using two separate methods. Overall, the best prediction by abiotic surrogates explained less than 30% of the pattern of biological similarity. Errors of false homogeneity were between 20% and 62%, depending on the methods of estimation. Predictive capability of abiotic surrogates at the taxon level was poor, with only 6% of taxon/surrogate correlations significant. Abiotic variables did not discriminate sufficiently between different soft bottom communities to be a reliable basis for mapping. These results have implications for the widespread use of abiotic surrogates in marine habitat mapping to plan for, or assess, representation in marine protected areas. Little confidence can be placed in marine habitat classifications based solely or largely on abiotic surrogates without calibration by rigorous biological surveys at the appropriate scale. Therefore, it is questionable whether marine protected areas designed on this basis can have measurable benefits for conservation.     

Impacts of a Hawaiian marine protected area network on the abundance and fishery sustainability of the yellow tang, Zebrasoma flavescens

Marine protected areas can enhance fish stocks within their boundaries, but the circumstances in which they might also supplement stocks or enhance fisheries outside their boundaries are less clear. Using visual survey and fishery data, we assess the impacts of increasing fishing effort, and of the establishment in Hawaii of a network of areas closed to aquarium fishing, on the prime-target species, yellow tang (Zebrasoma flavescens), and draw conclusions about MPA impacts on long-term fishery sustainability. Between 1999, when 27.8% of the coastline was closed to collecting, and 2007, the number of active fishers and total catch of yellow tang doubled. Prior to MPA establishment, yellow tang densities were similar at sites open to fishing and those slated for closure. By 2007, closed areas had five times the density of prime targeted sized fish (5-10 cm), and 48% higher density of adults than open areas. Densities of adults in ‘boundary’ areas (open areas <1 km from nearest MPA boundary) were significantly higher than in open areas far from MPA boundaries, which was indicative of spillover at that scale. Given the long life-span of yellow tang (>40 years) relative to the duration of protection and the increasing intensity of fishing, the likelihood is that protected areas will become increasingly important sources for the adult fishes which will sustain stocks and the fishery over the longer term.     

When the species is also a habitat: Comparing the predictively modelled distributions of Lophelia pertusa and the reef habitat it forms

Internationally there is political momentum to establish networks of marine protected areas for the conservation of threatened species and habitats. Practical implementation of such networks requires an understanding of the distribution of these species and habitats. Predictive modelling provides a method by which continuous distribution maps can be produced from limited sample data. This method is particularly useful in the deep sea where a number of biological communities have been identified as vulnerable ‘habitats’, including Lophelia pertusa reefs. Recent modelling efforts have focused on predicting the distribution of this species. However the species is widely distributed where as reef habitat is not. This study uses Maxent predictive modelling to investigate whether the distribution of the species acts as a suitable proxy for the reef habitat. Models of both species and habitat distribution across Hatton Bank and George Bligh Bank are constructed using multibeam bathymetry, interpreted substrate and geomorphology layers, and derived layers of bathymetric position index (BPI), rugosity, slope and aspect. Species and reef presence records were obtained from video observations. For both models performance is fair to excellent assessed using AUC and additional threshold dependant metrics. 7.17% of the study area is predicted as highly suitable for the species presence while only 0.56% is suitable for reef presence, using the sensitivity–specificity sum maximisation approach to determine the appropriate threshold. Substrate is the most important variable in the both models followed by geomorphology in the RD model and fine scale BPI in the SD model. The difference in the distributions of reef and species suggest that mapping efforts should focus on the habitat rather than the species at fine (100 m) scales.     

Long-term trends in lobster populations in a partially protected vs. no-take Marine Park

Increasing the level of protection afforded to the marine environment requires assessment of the efficacy of existing marine protected areas (MPAs) in protecting exploited species. Long-term data from before and after the establishment of MPAs provide a rare but valuable opportunity to assess these effects. In this study we present long-term data (1977-2005) from before and after park establishment, on the abundance of spiny lobster Jasus edwardsii from fixed sites in a no-take marine park and a recreationally fished marine park, to assess the efficacy of no-take vs. partial protection. Lobster densities were comparable between both marine parks prior to park establishment, but the response of lobster populations differed markedly following protection. On average, legal-sized lobster were eleven times more abundant and biomass 25 times higher in the no-take marine park following park establishment, while in the partially protected marine park there has been no significant change in lobster numbers. Furthermore, no difference was found in densities of legal-sized lobster between the partially protected marine park and nearby fully-fished sites (<1 per 500 m²). Long-term data from fully fished and partially protected sites suggest long-term declines in lobster populations and reflect regional patterns in catch per unit effort estimates for the fishery. The long-term patterns presented provide an unequivocal example of the recovery of lobster populations in no-take MPAs, but clearly demonstrate that allowing recreational fishing in MPAs has little benefit to populations of exploited species such as J. edwardsii.     

Ecosystem consideration in conservation planning: energy demand of foraging bottlenose whales (Hyperoodon ampullatus) in a marine protected area

The Gully, a submarine canyon off eastern Canada, was nominated as a pilot Marine Protected Area (MPA) in 1998, largely to safeguard the vulnerable population of northern bottlenose whales (Hyperoodon ampullatus) found there. The boundaries and ultimate management regime for the MPA for this area remain under review. We have estimated the energy consumption of bottlenose whales in the Gully based on the number of whales present at any time, their trophic level, the food requirements of each whale, and the rates of energy transfer between trophic levels. These calculations suggest that there must be a substantial spatial subsidy in the underlying foodweb of the submarine canyon to support the bottlenose whales using the Gully. A substantial area beyond the distribution of bottlenose whales in the area will therefore require protection. Conservation priorities to protect such subsidies will primarily involve additional protection at the level of the sea floor. Spatial subsidies are probably common in the marine environment, urging careful ecological analysis in the establishment of marine reserves and suggesting that conservation priorities need to take into account key ecological linkages and processes that are vital for sustaining species and habitats of concern.     

The impact of marine reserves on nekton diversity and community composition in subtropical eastern Australia

The effectiveness of marine reserve protection on the biodiversity of aquatic assemblages (i.e. nekton) in subtropical eastern Australia was examined within two small (<6 km²) marine reserves and four non-reserve areas. The two marine reserves, and their corresponding non-reserves, were located in different geographical locations within Moreton Bay (north and south) and sites were surveyed with multiple hauls of a seine net. Species richness, evenness, density and mean size of the inshore communities were compared between the reserves and non-reserves. No statistical significant difference was detected in species richness between the areas however species evenness was significantly lower in the only non-reserve site impacted by commercial net fishing. Mean size of nekton was found to be significantly greater in the marine reserves compared to non-reserves but no statistical significant difference was found in the density of nekton between the study sites. Multivariate analysis revealed differences in community composition, particularly between the geographical locations where areas were impacted by different types of fishing pressure (recreational v commercial). These results highlight the impact commercial fishing can have on entire nekton assemblages, not just on targeted species. Our study demonstrates that the small marine reserves in Moreton Bay are protecting marine biodiversity and are thus at least partially achieving their management objective (to enhance the zone’s marine biodiversity).     

Community-based conservation results in the recovery of reef fish spawning aggregations in the Coral Triangle

In the Coral Triangle community-based marine protected areas (MPAs) are being established at a prolific rate. Their establishment can benefit both fisheries and biodiversity, and they provide both a socially and economically acceptable means of managing coral reefs in developing nations. However, because such MPAs are typically small (usually <0.5 km²), they will rarely provide protection to large mobile fishes. An exception to this limitation may exist when community-based MPAs are established to protect small sites where vital processes occur, such as fish spawning aggregations (FSAs). To test the effectiveness of small (0.1–0.2 km²) MPAs for protecting FSAs, we monitored three FSA sites where brown-marbled grouper (Epinephelus fuscoguttatus), camouflage grouper (Epinephelus polyphekadion) and squaretail coralgrouper (Plectropomus areolatus) aggregate to spawn. Sites were monitored during peak reproductive periods (several days prior to each new moon) between January 2005 and November 2009. All three sites are located in New Ireland Province, Papua New Guinea, and had been exploited for decades, but in 2004 two sites were protected by the establishment of community-based MPAs. The third site continued to be exploited. Over the monitoring period densities of E. fuscoguttatus and E. polyphekadion increased at both MPAs, but not at the site that remained open to fishing. At one MPA the densities of E. polyphekadion increased tenfold. Our findings demonstrate that community-based MPAs that are appropriately designed and adequately enforced can lead to the recovery of populations of vulnerable species that aggregate to spawn.     

Entanglement of monofilament fishing lines and coral death

This study examined the extent of damage by monofilament fishing lines on the cauliflower coral (Pocillopora meandrina) colonies at a popular cast fishing site in Oahu, Hawaii in March 1998. Sixty-five percent of coral colonies had fishing lines on their surface, and 80% of colonies were either entirely or partially dead. The percentage of entirely and partially dead colonies was positively correlated with the percentage of colonies entangled with fishing lines. Percent dead coral surface area was larger for colonies with fishing lines than colonies without fishing lines. In colonies with fishing lines, percent dead coral surface area was positively correlated with percent area with fishing lines. The high incidence of coral colonies with fishing lines and the three sets of significant results indicated destructive effects of monofilament fishing lines to P. meandrina corals. There is thus a clear case that reefs should be protected not only from the effects of excessive removal of fishes, but also from the effects of unsuccessful fishing that results in fishing-line entanglement on corals.     

Effects of fishing and protection on Brazilian reef fishes

The vast Brazilian coast harbors unique and diverse reef fish communities. Unfortunately, relatively little is known about the impact of fishing on these fish species, and few management or conservation efforts are being made to protect them. Here, we examine the effect of different levels of protection on the composition, abundance, and size structure of reef fish species along a 2500 km portion of the Brazilian coastline, noting in particular the relative abundance of endemics and the effect of protection on these species. Pairwise comparisons of sites with different protection status (more versus less protected) were used to determine the potential responses of reef fishes to the establishment of marine protected areas. Highly targeted species (top predators and large herbivores) were significantly more abundant and larger in size within sites with a higher degree of protection, indicating that they benefit from protection, while lightly fished and unfished species were not. These results are consistent with past work documenting the responses of species to protection. Here, we use our results in particular to suggest strategies and provide expectations for managing and protecting Brazilian reef fisheries. Because this biogeographic province lies entirely within the jurisdiction of a single nation, there may be unique and significant opportunities to effectively manage and conserve these fish species.     

Assessing the effects of marine protected area (MPA) on a reef fish assemblage in a northwestern Mediterranean marine reserve: Identifying community-based indicators

Marine protected areas (MPAs) are increasingly envisaged as a tool to manage coastal ecosystems and fisheries. Assessment of their performance with respect to management objectives is therefore important. A number of MPAs provided conservation benefits for fished species. Observed benefits do not apply to all species at all times, and responses to protection are also highly variable among fish taxa. Among the many empirical studies on marine reserves, only a few designs considered ‘before and after data’ and spatial variation. In this paper, we are interested in assessing the effect of a no-take reserve on the reef fish assemblage in a northwestern Mediterranean example. Data were obtained from a three-year survey using underwater visual censuses (UVC), before and after MPA establishment. Permutational multivariate analysis of variance (PERMANOVA) and multivariate regression trees (MRT) were used to evaluate the effects of reserve protection on the reef fish assemblage, while accounting for habitat. Modelled biological responses were abundances and diversity indices calculated at different levels of the assemblage. Significant effects were found for many of these metrics. In addition to PERMANOVA, univariate models provided more insight into the magnitude and direction of effects. The most sensitive metrics were related to large species and species targeted by fishing. These results may be used to choose the metrics that are more suitable as community-based indicators of MPA impact in the perspective of monitoring programs.