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.     

Response of a desert lizard community to habitat degradation: Do ideas about habitat specialists/generalists hold?

We report the impact of human-induced desertification on the species richness, abundance, and composition of sand dune flora and herpetofauna of North Sinai, Egypt. Our hypothesis was that degraded habitats would have reduced vegetation complexity, richness, and abundance, and consequently lower reptile species richness and abundance. We also hypothesized that desert lizards would not follow the typical generalist/specialist responses to habitat degradation found in other biomes. Instead, we predicted that because vegetation loss intensifies the environmental extremity of deserts, those species specialized for open and sandy environments would be more likely to persist in desertified habitats than would desert generalists. Our results showed that areas protected from vegetation loss did not have significantly higher vegetation richness or abundance except for only a few species. However, protected sites did have significantly higher percent vegetation cover and height. Habitat protection clearly had strong effects on the reptile community as species richness and abundances were significantly higher in protected sites. The composition of the reptile community between protected and unprotected sites differed significantly. Contrary to past studies in other environments, desert generalist species were not able to persist in degraded sites and were only found in protected sites. Specialist species were ubiquitous in that they occurred in both areas protected and unprotected from vegetation loss. We propose that the effects of disturbance on species composition (specialists or generalists) depends on whether the disturbance exacerbates or reduces environmental harshness and the conditions that favor specialization. In extreme environments, specialist and generalist responses to habitat degradation are opposite to that of more productive environments.     

Small marine reserves can offer long term protection to an endangered fish

The protection of exploited species offered by marine reserves is maximised if individual fish live within the boundaries of the reserve for most of their lives. The direct benefit of this protection, the so called ‘reserve effect’, is the build-up of biomass through an increase in individual numbers and size. This effect can take several years depending on the life history traits of the target species. However, empirical studies on individual fish spatial behaviour at temporal scales long enough to be compatible with that of invoking the reserve effect are still virtually lacking. We studied the inter-annual patterns of habitat use of the dusky grouper (Epinephelusmarginatus), an endangered reef fish predator, using passive acoustic telemetry inside and around a small marine reserve in the Azores, mid-north Atlantic. All 11 individuals resided in the reserve year-round for up to 5 years. During these periods, fish were detected nearly every day and utilised stable core activity areas. Two smaller fish apparently abandoned the area after some time, probably relocating outside the reserve. Our results indicate that even small marine reserves can promote the long-term recovery of some endangered species if the long-term behaviour (i.e. site fidelity) of individuals is appropriate. This result bears direct implications for most existing coastal marine reserves, given that they fall into the small size range and many overexploited reef fish species may be highly site attached.     

Effects of fishing pressure and trophic group on abundance and spillover across boundaries of a no-take zone

We investigated factors influencing fish abundance and emigration across the boundaries of a no-take zone (NTZ), and its adjacent fished zones, within the Nabq Managed Resource Protected Area, South Sinai, Egyptian Red Sea. Underwater visual census of eight families of reef fish was undertaken at three depths across the whole NTZ (1.2 km), and for a similar distance into the adjacent fished zones. Because most fishing occurs in shallow water, the effect of the NTZ changed with depth. On reef flat transects (1 m depth), seven families showed significantly higher abundance in the NTZ, whereas at 10 m depth only two herbivorous families, Acanthuridae (surgeonfishes) and Siganidae (rabbitfishes) showed a significant difference, both being more abundant in the fished zones, an effect most likely due to reduced competition or predation. To look for evidence of emigration (spillover) of fish from the NTZ, data were also tested for evidence of gradients in abundance extending from within the NTZ to within the fished zones. In shallow water six families showed significant declines in abundance moving away from the centre of the NTZ, whereas at 10 m, the only significant gradients were for acanthurids and siganids, both becoming more abundant moving further into the fished zones. Comparisons of estimated fish length between areas also showed significant differences. The results indicate that at Nabq, while fishing reduces the abundance of most families, especially predators, abundance of some families and species, especially herbivores, may increase under light fishing pressure. Thus, spillover may occur to an extent and in a direction depending on trophic group and fishing intensity.     

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).     

Effectiveness of five small Philippines’ coral reef reserves for fish populations depends on site-specific factors, particularly enforcement history

Community-managed, no-take marine reserves are increasingly promoted as a simple, precautionary measure to conserve biodiversity and sustain coral reef fisheries. However, we need to demonstrate the effects of such reserves to those affected by the loss of potential fishing grounds and the wider scientific community. We surveyed changes in fish communities in five small marine reserves in the central Philippines and three distant Control sites over seven years. We conducted underwater visual censuses of 53 fish families within the reserve (Inside), with a kilometre of the boundary (Outside) and at Control sites. We found significant differences between fish communities Inside and Outside the reserve only at the two sites with strictest compliance with fishing prohibition, while there were significant differences to distant Control sites in all cases. The strongest responses to reserve protection were found in predatory fishes (groupers and breams) and in butterflyfish. Other abundant fish families showed weak effects of protection. For all taxa analysed, we found significant effects of reserve Site and Site × Treatment interactions. The detection of fish responses to reserves is complicated by potential spillover effects, site-specific factors, particularly compliance, and the difficulty of identifying appropriate control areas.     

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.     

Anthropogenic determinants of primate and carnivore local extinctions in a fragmented forest landscape of southern Amazonia

Habitat fragmentation has been shown to influence the abundance, movements and persistence of many species. Here, we examine the effects of forest patch and landscape metrics, and levels of forest disturbance on the patterns of local extinction of five primate and 14 carnivore species within 129 forest patches in a highly fragmented forest landscape of southern Brazilian Amazonia. Classic habitat area effects were the strongest predictors of species persistence, explaining between 42% and 55% of the overall variation in primate and carnivore species richness. Logistic regression models showed that anthropogenic disturbance, including surface wildfires, timber extraction and hunting pressure, had detrimental effects on the persistence of some species over and above those of fragment size. Different species ranged in their responses from highly sensitive to highly tolerant to forest fragmentation. Patterns of local extinction documented here were by no means chance events, and the nestedness of the overall species-by-site matrix was highly nonrandom in terms of the sets of species extirpated from the most to the least species-rich forest patches.     

Improving social acceptability of marine protected area networks: A method for estimating opportunity costs to multiple gear types in both fished and currently unfished areas

We present a novel method for calculating the opportunity costs to fishers from their displacement by the establishment of marine protected areas (MPAs). We used a fishing community in Kubulau District, Fiji to demonstrate this method. We modelled opportunity costs as a function of food fish abundance and probability of catch, based on gear type and market value of species. Count models (including Poisson, negative binomial and two zero-inflated models) were used to predict spatial abundance of preferred target fish species and were validated against field surveys. A profit model was used to investigate the effect of restricted access to transport on costs to fishers. Spatial distributions of fish within the three most frequently sighted food fish families (Acanthuridae, Lutjanidae, Scaridae) varied, with greatest densities of Lutjanidae and Acanthuridae on barrier forereefs and greatest densities of Scaridae on submerged reefs. Modelled opportunity cost indicated that highest costs to fishers arise from restricting access to the barrier forereefs. We included our opportunity cost model in Marxan, a decision support tool used for MPA design, to examine potential MPA configurations for Kubulau District, Fiji Islands. We identified optimum areas for protection in Kubulau with: (a) the current MPA network locked in place; and (b) a clean-slate approach. Our method of modelling opportunity cost gives an unbiased estimate for multiple gear types in a marine environment and can be applied to other regions using existing species data.     

Just passing through: Global change and the conservation of biodiversity in protected areas

Climate and land-use changes are expected to cause many species to shift into or beyond the boundaries of protected areas, leading to large turnover in species composition. Here, we tested whether long-established protected areas in Canada were more robust to such climate change impacts than areas with no formal protection by measuring changes in modeled butterfly species distributions (n = 139) within them. We used a recently established distribution modeling technique, Maxent, to model butterfly species’ distributions in two epochs (1900-1930 and 1960-1990). We compared rates of butterfly species richness and composition change within protected areas against distributions of randomly selected, ecologically similar, but non-protected, areas. Change in species richness and composition within protected areas were, for the most part, the same as changes observed among random areas outside protected area boundaries. These results suggest that existing protected area networks in Canada have provided little buffer against the impacts of climate change on butterfly species richness, possibly because land-use change surrounding long-standing protected areas has not been substantial enough to elevate the habitat protection afforded by these protected areas relative to other areas. Although protected areas are unarguably beneficial in conserving biological diversity, their capacity to maintain habitat appears insufficient to prevent broader-scale climate changes from sweeping species beyond their boundaries.     

Do we protect biological originality in protected areas? A new index and an application to the Bonifacio Strait Natural Reserve

Changes in biodiversity may disrupt the ecological functions performed by species assemblages. Hence, we urgently need to examine the implications of biodiversity loss not only in terms of species conservation but also in terms of sustainability of ecosystem services. The ability of protected areas to maintain local species richness has been clearly demonstrated. However, preserving goods and services provided by ecosystems requires not only the conservation of species richness but also the conservation of the most ‘original’ species, i.e. the ones with the highest average rarity of their attributes which are likely to perform some unique functions in ecosystems. We proposed a new conservation of biological originality (CBO) index as well as associated randomization tests to quantify the ability of protected areas to maintain viable populations for the most original species. As an application, we used long-term fisheries data collected in the Bonifacio Strait Natural Reserve (BSNR) to determine the species which benefited from the protection reinforcement in 1999. We also estimated a set of 14 ecomorphological functional traits on the 37 fish species caught in the BSNR and we obtained a functional originality value for each species. As a result, we found that functional originality was significantly protected in the fish assemblage of the BSNR: species with the most original functional trait combinations became more abundant after 1999. Our finding suggests that protecting most original species is an insurance against functional diversity erosion in the BSNR. More generally, our new index can be used to test whether protected areas may protect preferentially the most original species and whether restorative management promotes the reestablishment of the most original species with particular habitat requirements.     

Effects of habitat characteristics and sedimentation on performance of marine reserves in St. Lucia

This study examines factors affecting the rate and extent of biomass build-up among commercially important groupers, snappers, grunts, parrotfish and surgeonfish in a network of four marine reserves in southwest St. Lucia, Caribbean. Reserves constituted 35% of the total reef area originally available for fishing. Protection was instigated in 1995 after a baseline survey with annual or biennial censuses performed until 2002. Each survey consisted of 114 fifteen minute fish counts in reserves and 83 in fishing grounds, at depths of 5 m and 15 m in a 10 m diameter counting area. Estimates of number and size (cm) of target species were used to calculate fish family biomass. Data were analysed using three-way ANOVA in a before-after-control-impact pairs (BACIP) design. All families increased significantly in biomass over time at nearly all sites. Increases were greater in reserves than fishing grounds, except for grunts, and responses were strongest in parrotfish and surgeonfish. The combined biomass of families more than quadrupled in reserves and tripled in fishing grounds between 1995 and 2002. During this period coral cover declined by 46% in reserves and 35% in fishing grounds. Multiple regression showed that neither habitat characteristics nor habitat deterioration significantly affected rates of biomass build-up. The key factor was protection from fishing, which explained 44% of the variance in biomass growth. A further 28% of the variance was explained by sedimentation, a process known to stress reef invertebrates, significantly reducing the rate of biomass build-up. St. Lucia’s reserves succeeded in producing significant gains to fish stocks despite coral cover and structural complexity falling steeply over the period of the study.     

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.     

Inter-annual variability in the home range of breeding turtles: Implications for current and future conservation management

We assessed home range size for breeding loggerhead turtles (Caretta caretta) near the limit of the species range at the Greek island of Zakynthos in the Mediterranean. Thirteen adult females and seven adult males were tracked using GPS units (loggers and transmitters) during May and June of 2006, 2007 and 2008. Kernel analysis indicated that core home range sizes (50% estimator; range: 2.9-19.7 km²) for both males and females were restricted to a 7.5 km tract of coastline. 15% of GPS locations fell outside of the national park protection zones, while within the protected breeding area 88% of GPS locations occurred in zones of minimal protection. Female home ranges were 64% larger in 2008 than in 2006 and 2007, indicating that several years monitoring may be required for the most effective designation of marine protected areas (MPAs). Ten of the tracked females departed the core breeding area on 15 occasions for periods of 1-15 days travelling distances of 10-100 km, although none nested at alternative breeding sites. The inter-annual variability of breeding area home range size and likelihood of incidence of forays appeared be correlated with barometric pressure. The movement responses of loggerheads to environmental conditions implicates an ability to switch nesting areas over small scales in response to climate change. However, such behaviour suggests the protection of existing core breeding sites may be inadequate, with policy makers being required to consider the protection of broader areas to encompass potential changes in the habitat needs of this species.     

The quest for cryptic creatures: Impacts of species-focused recreational diving on corals

Coral reefs are popular with ecotourists, but the impact of divers on reefs is cause for concern. In this study, we assessed the damage to corals caused by divers seeking cryptic but charismatic fish such as seahorses (family Syngnathidae) and frogfishes (Antennariidae), which are found on reefs around the world. These fish are closely associated with the reef substratum, thus bringing divers into close proximity to coral. We found that when in the vicinity of frogfish and seahorses, divers made unintentional contact with corals significantly more often and for longer periods than when these species were absent. This change in diver behaviour resulted in a greater frequency of coral breakage and scarring at seahorse/frogfish sites than at ecologically equivalent control sites. However, the spatial extent of damage appeared limited. Beyond 1-3 m from the seahorse or frogfish, coral breakage and scarring rate decreased to levels similar to those of control sites. None of the coral species, which suffered the most damage, was particularly rare, suggesting that the habitat conservation concern of these marked shifts in diver behaviour is limited. Nevertheless, the use of pre-dive briefings and smaller dive group sizes could minimise the damage caused by divers approaching cryptic species of interest near the reef.     

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 effects of forest patch size and matrix type on changes in carabid beetle assemblages in an urbanized landscape

In this study we compared ground beetles (Carabidae) from a range of different forest fragments along an urbanization gradient in Brussels, Belgium. We address the following questions: (i) How does the degree of urbanization in the surrounding habitat affect forest beetles, and does it interact with the effects of patch size and distance to forest edge? (ii) Do these factors have a different effect at the level of individual species, habitat affinity groups or total community? During 2002 we sampled 13 forest plots in 10 forest patches, ranging in size from 5.27 to 4383 ha. The beetles were captured using transects of pitfall traps from the edge to a distance of 100 m into each woodland and identified to species level. Effects of urbanization, forest size and forest edge were evaluated on total species number, abundance and habitat affinity groups and ten abundant, widespread model carabid species. Overall, the effects of urbanization, forest size and edge effects slightly influenced total species richness and abundance but appeared to have a major effect on ground beetle assemblages through species specific responses. More urbanized sites had significantly fewer forest specialists and more generalist species. Large forest fragments were favoured by forest specialist species while generalist species and species frequently associated with forest (forest generalists) dominated the smaller forests. Forest edges mainly harboured generalist species while forest specialist species were more frequent into the forests if the forest patches were large enough, otherwise they disappeared due to the destruction or impoverishment of their habitat. Our results show the importance of differentiating between habitat affinity, especially habitat generalists versus specialists, the latter having a higher value in nature conservation, and merely the quantity of species represented in human-dominated areas.     

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.     

Relationship between distributions of threatened plants and protected areas in Britain

The establishment and maintenance of a system of protected areas is central to regional and global strategies for the conservation of biodiversity. The current global trend towards human population growth and widespread environmental degradation means that such areas are becoming increasingly isolated in fragmented habitat islands. In regions in which this process is well advanced a high proportion of species are thus predicted to have become restricted to protected areas. Here using uniquely detailed datasets for Britain, a region with close to the global level of percentage coverage by statutory protected areas, we determine the extent of restriction of Red List vascular plant species of conservation concern to these areas. On the basis of currently known distributions, overall our results strongly support the importance of a dual conservation strategy in Britain, in which protected areas are maintained with particular reference to those biodiversity features (such as many threatened plant species) that are highly dependent on them, and in which components of the wider landscape are also managed in such a way as to promote the abundance and distribution of such features with particular reference to those which are unlikely to persist in protected areas alone.     

Gradients of abundance and biomass across reserve boundaries in six Mediterranean marine protected areas: Evidence of fish spillover?

Marine protected areas (MPAs) are considered as an effective tool in marine coastal management, and considered able to enhance local fisheries through adult fish spillover. Indirect evidence of fish spillover could be obtained by horizontal gradients in fish abundance. To address this question, the existence of gradients of fish abundance and biomass across marine reserve boundaries was assessed in six Mediterranean MPAs using underwater visual censuses performed at various distances from the core of the MPA, in integral reserve (IR), to buffer zone (BZ) and fished areas. A reserve effect was evidenced with higher values of fish species richness (×1.1), abundance (×1.3), and biomass (×4.7) recorded inside MPAs compared to adjacent fished areas. Linear correlations revealed significant negative gradients in mean fish biomass in all the reserves studied after the effect of habitat had been removed, whereas negative gradients in abundance were less conspicuous. Generalized additive models suggested two main patterns of biomass gradients, with a sharp decrease at the IR-BZ boundary or at the BZ-fished area boundary. It was estimated that fish spillover beneficial to local fisheries occurred mostly at a small spatial scale (100s of metres). The existence of regular patterns of negative fish biomass gradients from within MPAs to fished areas was consistent with the hypothesis of adult fish biomass spillover processes from marine reserves and could be considered as a general pattern in this Mediterranean region.