Marine protected areas export larvae of infauna, but not of bioengineering mussels to adjacent areas

Populations of organisms that create habitat can often be fragmented throughout landscapes by anthropogenic disturbances such as harvesting and loss or change to the identity of such bioengineers may lead to large changes in biodiversity. Using the fauna associated with a bioengineer, the intertidal mussel Perna perna, we tested hypotheses about the relative importance of larval export from protected populations in marine reserves. Harvesting led to the replacement of P. perna and the domination of shores outside reserves by turf-forming coralline algae, mostly Corallina spp. We determined whether the diverse fauna recruiting onto artificial units of habitat placed within mussel beds differed between reserves and non-reserve areas or whether shores outside reserves, and open to harvesting, received recruits through larval export from reserves. Furthermore, we determined whether this was affected by the distance away from reserves and whether colonisation was achieved by movement of adults from surrounding biogenic habitats or via the plankton. Overall, we found no effect of increasing distance away from a reserve on the cover of adult mussels or associated fauna. We found strong effects of the presence of marine reserves on abundances of molluscs and polychaetes but not crustaceans. There were greater densities of molluscs in sites with a reserve (i.e. inside reserves, and up to 5 km outside reserve boundaries), but more polychaetes in exploited sites. For molluscs, this pattern was driven by gastropods rather than bivalves. Furthermore, although reserves had greater cover of adult mussels than non-reserve areas, recruitment of mussels was not greater inside or near to reserves. Our study illustrates the effectiveness of these reserves in protecting stocks of adult mussels, and although there was no evidence that reserves provided export of the larvae of mussels (the target species), they did provide larval export of non-targeted associated species. By protecting a harvested bioengineer and through export of the larvae of its associated fauna, these reserves fulfil some, but not all the conservation aims of a marine protected area.     

Interactions between poaching and management policy affect marine reserves as conservation tools

To explore the effects of poaching within marine reserve boundaries under three different management policies this analysis uses a simple age-structured reserve model based on yield maximization or reproductive thresholds of Black rockfish (Sebastes melanops). Departures from the traditional assumptions of full compliance to reserve boundaries alter the conclusions of prior modeling work that demonstrate yield equivalence to no-reserve effort control management and augmented reproductive benefits when small reserves are implemented. By degrading the recruitment subsidization effect to nonreserve areas from protected reserve populations, poaching resulted in negative externalities for compliant fishermen in open areas in terms of yield and degraded the reproductive output and age-structure of the system. All three policies required effort reduction in open areas as a response to poaching in reserves. The strength of the impacts from poaching varied with policy choice and harvest intensity in the reserve, where at the highest level of poaching modeled here (15% annual exploitation rate of the vulnerable reserve population) biological and fishery benefits of implementing reserves were totally negated. Under the assumptions of this model, a policy managing for a reproductive threshold that excludes the reserve population is the precautionary choice if poaching is likely. The results of this exercise emphasize the importance of garnering compliance to reserve boundaries from resource-users for spatial closures to be successful ocean management tools.     

Designing marine reserves for interacting species: Insights from theory

The primary goals of marine reserves include protecting biodiversity and ecosystem structure. Therefore, a multispecies approach to designing and monitoring reserve networks is necessary. To gain insight into how the interactions between species in marine communities may affect reserve design, we synthesize marine reserve community models and community models with habitat destruction and fragmentation, and we develop new extensions of existing models. This synthesis highlights the potential for species interactions to alter reserve design criteria; in particular, accounting for species interactions often leads to an increase in reserve size necessary to protect populations. Accounting for species interactions also indicates the need to base reserve design and monitoring on a variety of species, especially long-distance dispersers, inferior colonizers, and specialists. Finally, the new model extensions highlight how, given dispersal, source populations outside reserves as well as increases in fished populations after reserve establishment may negatively affect reserve populations of competitors or prey. Therefore, multispecies harvest dynamics outside reserves and before reserve establishment are critical to determining the appropriate reserve size, spacing, and expectations after establishment. These models highlight the importance of species interactions to reserve design and provide guidelines for how this complexity can begin to be incorporated into conservation planning.     

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).     

Spatial assessment of benefits of a coastal Mediterranean Marine Protected Area

Marine reserves are increasingly advocated not only as conservation but also as fisheries management tools to safeguard the decline of coastal fishing resources. Still, conclusive evidence of their functioning is lacking, amongst others due to the influence of spatio-temporal variations in fish populations and habitat heterogeneity which could hamper a sound data interpretation. We conducted a spatial analysis of the benefits of the Medes Island Marine Reserve by combining geostatistical and Geographic Information System (GIS) tools. Concurrently, we analysed effects of trends reflecting habitat heterogeneity and spatial structuring of data on spatial predictions of fish catch per unit effort (CPUE) and length. Predicted spatial patterns showed the complexity and simultaneous action of trend factors leading to mostly non-linear gradients in CPUE and length data. CPUE of total fish and CPUE and length of common pandora (Pagellus erythrinus) increased close to the Integral Reserve due to direct and indirect reserve effects. CPUE and length of striped red mullet (Mullus surmuletus) slightly increased also near the Integral Reserve, but distinct reserve effects could not be identified due to the strong influence of artificial reefs. We conclude that the spatial dimension of the Buffer Zone, where artisanal fisheries are allowed, permits in general protection only for target species, favouring a habitat with no discontinuities from the reserve outwards. Our spatial approach to assess reserve benefits provides major insights into complex systems like coastal marine reserves in the northwestern Mediterranean. In addition, it contributes to a crucial aspect of marine conservation, viz. the decision on the spatial dimension of protected areas.     

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.     

The effects of including marine ecological values in terrestrial reserve planning for a forest-nesting seabird

Theoretical advances in systematic reserve design aim to promote the efficient use of limited conservation resources and to increase the likelihood that reserve networks enhance the persistence of valued species and ecosystems. However, these methods have rarely been applied to species that rely on spatially disjunct habitats. We used the marbled murrelet, a seabird that requires old-growth forest in which to nest and high quality marine habitats in which to forage, as a case study to explore methods of incorporating multiple ecological values into single species spatial reserve design. Specifically, we used the cost function in MARXAN to include the ecological value of marine habitats while identifying spatial solutions for terrestrial nesting habitat reserves. Including marine values influenced terrestrial reserve designs most when terrestrial habitat targets were low and little or none of the target was represented in pre-existing protected areas. Our results suggest that including marine values in the planning process will influence marbled murrelet terrestrial reserve designs most where substantial terrestrial nesting habitat still exists, where new reserves are relatively unconstrained by pre-existing reserves, or when conservation resources only allow the protection of a small fraction of available habitat. This paper presents a novel framework for incorporating multiple measures of ecological value in the spatial reserve design process and should be particularly useful for species that rely on multiple habitats during their life cycle.     

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.     

Consequences of adult and juvenile movement for marine protected areas

Adult and juvenile mobility has a considerable influence on the functioning of marine protected areas. It is recognized that adult and juvenile movement reduces the core benefits of protected areas, namely protecting the full age–structure of marine populations, while at the same time perhaps improving fisheries yield over the no-reserve situation through export of individuals from protected areas. Nevertheless, the study of the consequences of movement on protected area functioning is unbalanced. Significant attention has been paid to the influence of certain movement patterns, such as diffusive movement and home ranges, while the impacts of others, such as density-dependent movements and ontogenetic migrations, have been relatively ignored. Here we review the diversity of density-independent and density-dependent movement patterns, as well as what is currently known about their consequences for the conservation and fisheries effects of marine protected areas. We highlight a number of ‘partially addressed’ issues in marine protected area research, such as the effects of reserves targeting specific life phases, and a number of essentially unstudied issues, such as density-dependent movements, nomadism, ontogenetic migrations, behavioral polymorphism and ‘dynamic’ reserves that adjust location as a realtime response to habitat changes. Assessing these issues will be essential to creating effective marine protected area networks for mobile species and accurately assessing reserve impacts on these species.     

Efficiency and effectiveness in representative reserve design in Canada: The contribution of existing protected areas

To be effective, reserve networks should represent all target species in protected areas that are large enough to ensure species persistence. Given limited resources to set aside protected areas for biodiversity conservation, and competing land uses, a prime consideration for the design of reserve networks is efficiency (the maximum biodiversity represented in a minimum number of sites). However, to be effective, networks may sacrifice efficiency. We used reserve selection algorithms to determine whether collections of existing individual protected areas in Canada were efficient and/or effective in terms of representing the diversity of disturbance-sensitive mammals in Canada in comparison to (1) an optimal network of reserves, and (2) sites selected at random. Unlike previous studies, we restricted our analysis to individual protected areas that met a criterion for minimum reserve size, to address issues of representation and persistence simultaneously. We also tested for effectiveness and efficiency using historical and present-day data to see whether protected area efficiency and/or effectiveness varied over time. In general, existing protected areas did not effectively capture the full suite of mammalian species diversity, nor are most existing protected areas part of a near-optimal solution set. To be effective, Canada’s network of reserves will require at minimum 22 additional areas of >2700 km². This study shows that even when only those reserves large enough to be effective are considered, protected areas systems may not be representative, nor were they representative at the time of establishment.     

Italian marine reserve effectiveness: Does enforcement matter?

Marine protected areas (MPAs) have become popular tools worldwide for ecosystem conservation and fishery management. Fish assemblages can benefit from protection provided by MPAs, especially those that include fully no-take reserves. Fish response to protection can thus be used to evaluate the effectiveness of marine reserves. Most target fish are high-level predators and their overfishing may affect entire communities through trophic cascades. In the Mediterranean rocky sublittoral, marine reserves may allow fish predators of sea urchins to recover and thus whole communities to be restored from coralline barrens to macroalgae. Such direct and indirect reserve effects, however, are likely to be related to the enforcement implemented. In Italy, many MPAs that include no-take reserves have been declared, but little effort has been spent to enforce them. This is a worldwide phenomenon (although more common in some regions than others) that may cause MPAs and reserves to fail to meet their targets. We found that 3 of 15 Italian marine reserves investigated had adequate enforcement, and that patterns of recovery of target fish were related to enforcement. No responses were detected when all reserves were analyzed as a whole, suggesting enforcement as an important factor to be considered in future studies particularly to avoid that positive ecological responses in properly managed reserves can be masked by neutral/negative results in paper parks. Positive responses were observed for large piscivores (e.g. dusky groupers) and sea urchin predators at reserves where enforcement was effective. Those reserves with low or null enforcement did not differ from fished areas.     

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.     

Assessing the performance of the existing and proposed network of marine protected areas to conserve marine biodiversity in Chile

The growing concern about the profound influence of human activities on marine ecosystems has been the driving force behind the creation of marine reserves in the last few decades. With almost 4200 km of coastline, Chile has not been the exception to this trend. A set of conservation priority sites has recently been proposed by the Chilean government to expand the current marine reserve network. In this study, we used the most comprehensive information currently available on the distribution of 2513 marine species in Chile to assess the efficiency of the existing system of marine protected areas (MPA) and the conservation priority sites identified by the government. Additionally, we evaluated the vulnerability of the reserve network selected with respect to threatening human activities. Our results show that both the existing protected areas and the proposed priority sites are relatively effective at protecting Chilean marine biodiversity. However, the majority of the species that are not represented within the existing or projected MPA network have very restricted distributions and are, therefore, of high conservation concern. To cover all species requires a network of 35 MPAs (46% of the total number of planning units). Many of the sites identified as irreplaceable present conflict with one or more human activities, particularly in the central region of the country. This study emphasizes the need for a systematic conservation planning approach to maximize the representation of species and prioritize those areas where conflicts between marine biodiversity conservation and human activities may occur.     

The island dilemma: Lessons of modern biogeographic studies for the design of natural reserves

A system of natural reserves, each surrounded by altered habitat, resembles a system of islands from the point of view of species restricted to natural habitats. Recent advances in island biogeography may provide a detailed basis for understanding what to expect of such a system of reserves. The main conclusions are as follows:The number of species that a reserve can hold at equilibrium is a function of its area and its isolation. Larger reserves, and reserves located close to other reserves, can hold more species.If most of the area of a habitat is destroyed, and a fraction of the area is saved as a reserve, the reserve will initially contain more species than it can hold at equilibrium. The excess will gradually go extinct. The smaller the reserve, the higher will be the extinction rates. Estimates of these extinction rates for bird and mammal species have recently become available in a few cases.Different species require different minimum areas to have a reasonable chance of survival.Some geometric design principles are suggested in order to optimise the function of reserves in saving species.     

Designing a network of marine reserves in the Mediterranean Sea with limited socio-economic data

The present study is the first to determine priorities for the location of marine reserves using spatial prioritization software in the eastern Mediterranean Sea. We used biophysical data from visual census surveys on: fish species abundance, presence of various habitat types, and percent coverage of seagrasses and canopy algae. Efficient conservation planning requires spatially explicit information on how proposed management will affect stakeholders, which in this region was very limited. We created novel socio-economic cost indices to account for fisheries and tourists. Our fishing metrics were based on fisher behaviour including information on the location of ports and areas often inaccessible to fishers due to high wind exposure. We developed a cost index for tourism based on the availability of beds for tourists. We examined how the spatial priorities for marine reserves varied using different combinations of these socio-economic cost metrics. We found about 17% of sites were a high priority regardless of which cost metric was used. We also compared, for the first time in the Mediterranean, our results devised using systematic conservation planning approaches with priorities developed by two non-systematic methods, the Natura 2000 proposed marine reserves and sites that local fishers proposed for protection. Only a few sites identified by our approach were the same as those recommended as part of Natura 2000 or the fishers’ proposals. This suggests that much more work is needed to harmonise existing proposals with the principles of efficient systematic conservation planning.     

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.     

Forest restitution and protected area effectiveness in post-socialist Romania

The effectiveness of protected areas can diminish during times of pronounced socio-economic and institutional change. Our goals were to assess the effectiveness of Romanian protected areas at stemming unsanctioned logging, and to assess post-socialist logging in their surrounding landscapes, during a time of massive socio-economic and institutional change. Our results suggest that forest cover remained fairly stable shortly before and after 1990, but forest disturbance rates increased sharply in two waves after 1995 and 2005. We found substantial disturbances inside protected areas, even within core reserve areas. Moreover, disturbances in the matrix surrounding protected areas were even lower than inside protected area boundaries. We suggest that these rates are largely the result of high logging rates, triggered by rapid ownership and institutional changes. These trends compromise the goals of Romania’s protected area network, lead to an increasing loss of forest habitat, and more isolated and more fragmented protected areas. The effectiveness of Romania’s protected area network in terms of its ability to safeguard biodiversity is therefore most likely decreasing.     

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.     

Responses to marine reserves: Decreased dispersion of the sparid Pagrus auratus (snapper)

Quantifying the movement of exploited species inside marine reserves is a popular research topic, yet few studies have quantified movement inside and outside of reserves. If individuals inside reserves behave differently to those outside, this information should be incorporated into reserve design and in management efforts to avoid the selective removal of certain behaviors. We used acoustic telemetry to monitor the movements of the sparid Pagrus auratus (snapper) inside and outside a marine reserve (the Leigh Marine Reserve, north-eastern New Zealand). We tagged 39 snapper within an array of acoustic receivers that encompassed reserve and fished areas. Nineteen snapper were resident over a 5-month period; the remainder either left the array or died. Residential fish expressed two home range types. One group had uni-modal home ranges that on average encompassed c.900 m linear distance. All nine residential snapper from the reserve displayed this behavior, as well as five of ten residential fish from the non-reserve area. The second group (five fish, all from the non-reserve area) had home ranges with two separate modes (bi-modal home ranges), which on average encompassed c.2 100 m linear distance. We suggest that some aspect of the marine reserve environment encourages extreme residency by either the modification of individual behaviors or through the removal of selective exploitation. If true this suggests that reserves and populations of exploited animals may become dependent on the life history characteristics of the individuals they encourage or select for.     

Bali Barat: An Indonesian marine protected area and its resources

Since the early 1970s the Indonesian Government has tried to establish marine nature reserves. The initial marine protected areas, however, have not been successful, due to lack of enforcement, and the present account is of a site on the western tip of Bali, where it is hoped that the idea will come to fruition. The marine resources of this Reserve, Bali Barat, are described, together with the various ways in which they are at present being exploited. The plan developed for the Reserve aims to prohibit destructive uses, while allowing those which are not as yet considered damaging to the area. The needs for strict protection of certain sites for tourism and for subsistence uses have been reconciled through a system of zones. The experience gained from Bali Barat should enable effective marine reserve conservation to be extended to other parts of the country, and the Reserve could also contribute directly to education of the public, and to research and management expertise.