Designing Effective Marine Protected Areas in Seaflower Biosphere Reserve, Colombia, Based on Biological and Sociological Information

Abstract:   Ecologists have paid increasing attention to the design of marine protected areas (MPAs), and their design advice consistently recommends representing all habitat types within MPAs or MPA networks as a means to provide protection to all parts of the natural ocean system. Recent developments of new habitat-mapping techniques make this advice more achievable, but the success of such an approach depends largely on our ability to define habitat types in a way that is ecologically relevant. We devised and tested the ecological relevance of a set of habitat-type definitions through our participation in a stakeholder-driven process to design a network of MPAs, focusing on no-take marine reserves in the Seaflower Biosphere Reserve, San Andrés Archipelago, Colombia. A priori definitions of habitat types were ecologically relevant, in that our habitat-type definitions corresponded to identifiable and unique characteristics in the ecological communities found there. The identification of ecological pathways and connectivity among habitats also helped in designing ecologically relevant reserve boundaries. Our findings contributed to the overall design process, along with our summary of other general principles of marine reserve design. Extensive stakeholder input provided information concerning the resources and their patterns of use. These inputs also contributed to the reserve design process. We anticipate success for the Seaflower Biosphere Reserve at achieving conservation and social goals because its zoning process includes detailed yet flexible scientific advice and the participation of stakeholders at every step .     

Long-term trends in invertebrate–habitat relationships under protected and fished conditions

Few studies examine the long-term effects of changing predator size and abundance on the habitat associations of resident organisms despite that this knowledge is critical to understand the ecosystem effects of fishing. Marine reserves offer the opportunity to determine ecosystem-level effects of manipulated predator densities, while parallel monitoring of adjacent fished areas allows separating these effects from regional-scale change. Relationships between two measures of benthic habitat structure (reef architecture and topographic complexity) and key invertebrate species were followed over 17 years at fished and protected subtidal rocky reefs associated with two southern Australian marine reserves. Two commercially harvested species, the southern rock lobster (Jasus edwardsii) and blacklip abalone (Haliotis rubra) were initially weakly associated with habitat structure across all fished and protected sites. The strength of association with habitat for both species increased markedly at protected sites 2 years after marine reserve declaration, and then gradually weakened over subsequent years. The increasing size of rock lobster within reserves apparently reduced their dependency on reef shelters as refuges from predation. Rising predation by fish and rock lobster in the reserves corresponded with weakening invertebrate–habitat relationships for H. rubra and sea urchins (Heliocidaris erythrogramma). These results emphasise that animal–habitat relationships are not necessarily stable through time and highlight the value of marine reserves as reference sites. Our work shows that fishery closures to enhance populations of commercially important and keystone species should be in areas with a range of habitat features to accommodate shifting ecological requirements with ontogenesis.     

Risks to large marine protected areas posed by drifting fish aggregation devices

Mapping and predicting the potential risk of fishing activities to large marine protected areas (MPAs), where management capacity is low but fish biomass may be globally important, is vital to prioritizing enforcement and maximizing conservation benefits. Drifting fish aggregating devices (dFADs) are a highly effective fishing method employed in purse seine fisheries that attract and accumulate biomass fish, making fish easier to catch. However, dFADs are associated with several negative impacts, including high bycatch rates and lost or abandoned dFADs becoming beached on sensitive coastal areas (e.g., coral reefs). Using Lagrangian particle modeling, we determined the potential transit of dFADs in a large MPA around the Chagos Archipelago in the central Indian Ocean. We then quantified the risk of dFADs beaching on the archipelago's reefs and atolls and determined the potential for dFADs to pass through the MPA, accumulate biomass while within, and export it into areas where it can be legally fished (i.e., transit). Over one-third (37.51%) of dFADs posed a risk of either beaching or transiting the MPA for >14 days, 17.70% posed a risk of beaching or transiting the MPA for >30 days, and 13.11% posed a risk of beaching or transiting the MPA for >40 days. Modeled dFADs deployed on the east and west of the perimeter were more likely to beach and have long transiting times (i.e., posed the highest risk). The Great Chagos Bank, the largest atoll in the archipelago, was the most likely site to be affected by dFADs beaching. Overall, understanding the interactions between static MPAs and drifting fishing gears is vital to developing suitable management plans to support enforcement of MPA boundaries and the functioning and sustainability of their associated biomass.     

Differential movement patterns and site fidelity among trophic groups of reef fishes in a Hawaiian marine protected area

We tracked the long-term movements of 70 parrotfishes, surgeonfishes and goatfishes captured inside a small (1.3 km²) marine protected area (MPA: Kealakekua Bay Marine Life Conservation District, Hawaii) by implanting them with small transmitters and deploying underwater monitoring devices inside the bay and along 100 km of the adjacent west Hawaii coastline. Individual fish were detected inside Kealakekua Bay for up to 612 days but many were detected for much shorter periods (median = 52 days). There were species-specific differences in the scale of movements and habitats used, but most fish utilized between 0.2 and 1.6 km of coastline, and individuals of each species showed some degree of diel habitat shift. A wide variety of reef fishes captured inside the MPA swam back and forth across an MPA boundary intersecting continuous reef (i.e., this boundary was porous to reef fish movements), but only 1 of 11 species tagged crossed a wide sandy channel inside Kealakekua Bay suggesting that this feature may function as a natural barrier to movements. Results indicate relatively small MPAs (<2 km of coastline) could provide effective, long-term protection for multi-species assemblages of reef fishes provided that boundaries are situated along major habitat breaks (e.g., large sand channels between reefs) that may serve as natural barriers to reef fish movements. It is crucial that a multi-species approach be used when assessing MPA effectiveness.     

Comanagement Practices Enhance Fisheries in Marine Protected Areas

Abstract:  Fishing activities worldwide have dramatically affected marine fish stocks and ecosystems. Marine protected areas (MPAs) with no-take zones may enhance fisheries, but empirical evidence of this is scant. We conducted a 4-year survey of fish catches around and within an MPA that was previously fully closed to fishing and then partially reopened under regulated comanaged fishing. In collaboration with the fishers and the MPA authority, we set the fishing effort and selected the gear to limit fishing impact on key fish predators, juvenile fish stage, and benthic communities and habitats. Within an adaptive comanagement framework, fishers agreed to reduce fishing effort if symptoms of overfishing were detected. We analyzed the temporal trends of catch per unit of effort (CPUE) of the whole species assemblages and CPUE of the four most valuable and frequent species observed inside the opened buffer zone and outside the MPA investigated. After the comanaged opening, CPUE first declined and then stabilized at levels more than twice that of catches obtained outside the MPA. Our results suggest that working closely with fishers can result in greater fisheries catches. Partial protection of coastal areas together with adaptive comanagement involving fishers, scientists, and managers can effectively achieve conservation and fishery management goals and benefit fishing communities and alleviate overfishing.     

Combined Effects of Levels of Protection and Environmental Variables at Different Spatial Resolutions on Fish Assemblages in a Marine Protected Area

Abstract: The links between species–environment relations and species’ responses to protection are unclear, but the objectives of marine protected areas (MPAs) are most likely to be achieved when those relations are known and inform MPA design. The components of a species’ habitat vary with the spatial resolution of the area considered. We characterized areas at two resolutions: 250 m² (transect) and approximately 30,000 m² (seascape). We considered three categories of environmental variables: substrate type, bottom complexity, and depth. We sought to determine at which resolution habitat characteristics were a better predictor of abundance and species composition of fishes and whether the relations with environmental variables at either resolution affected species’ responses to protection. Habitat features accounted for a larger proportion of spatial variation in species composition and abundances than differences in protection status. This spatial variation was explained best by habitat characteristics at the seascape level than at the transect level. Species’ responses to protected areas were specific to particular seascape characteristics, primarily depth, and bottom complexity. Our method may be useful for prioritizing marine areas for protection, designing MPAs, and monitoring their effectiveness. It identified areas that provided natural shelter, areas acting as buffer zones, and areas where fish species were most responsive to protection. The identification of such areas is necessary for cost‐effective establishment and monitoring of MPAs.     

Seasonal,diel, and tidal movements of green jobfish (<Emphasis Type="Italic">Aprion virescens</Emphasis>, Lutjanidae) at remote Hawaiian atolls: implications for marine protected area design

Empirical data quantifying the long-term movement patterns of coral reef top predators are needed in order to design marine protected areas (MPAs) that will provide these fishes with effective, long-term protection. Acoustic telemetry was used to quantify the movements of a large coral reef top predator (Aprion virescens, Lutjanidae; Hawaiian name ‘uku’) at five atolls in the Northwestern Hawaiian Islands Marine National Monument (NWHIMNM) from May 2005 to September 2006. The study atolls were located between 23.8°N, 166.2°W and 28.5°N, 178.3°W, and were separated from their nearest receiver-equipped neighbor by distances ranging from 100 to 478 km. No inter-atoll movements by uku were detected but individuals were seasonally site-attached to core activity areas of up to 12 km in length, and ranged up to 19 km across atolls. Within their core areas, tagged uku exhibited diel and tidal habitat shifts, with the latter resulting in round trips of up to 24 km in 24 h. Seasonal uku migrations resulted in extended winter (October–April) absences from summer (May–September) core activity areas and may be linked to summer spawning. Large MPAs (i.e., entire islands, atolls or banks) would probably be required for full protection of resident populations of adult uku, but such ‘island-scale’ MPAs will not benefit fisheries unless there is significant larval supply from MPAs to neighboring fished areas, or adult emigration over time scales exceeding the 16-month monitoring period of this study. A mixed management strategy of combining smaller MPAs with conventional measures (e.g., minimum size limits, catch, and effort restrictions) may be the best approach for sustaining uku fisheries. However, this would still require relatively large MPAs (12 km in length) to contain uku short-term (diel and tidal) movements, and seasonal migrations would still take uku beyond the boundaries of MPAs of this size. These fluxes across MPA boundaries could supply fish to fisheries but, if high exploitation rates exist, fishing could eliminate key MPA benefits such as increased numbers of large, highly fecund individuals. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Indirect Benefits of Marine Protected Areas for Juvenile Abalone

Abstract: Marine protected areas ( MPAs) designed to provide harvest refugia for red sea urchins (  Strongylocentrotus franciscanus ) offer a unique opportunity to study the indirect effects of urchin fishing on subtidal communities. Sea urchins may provide important cryptic microhabitat for juvenile abalone sheltering beneath urchin spines in shallow habitats worldwide. We investigated the abundance of juvenile (3–90 mm) red abalone, (    Haliotis rufescens ) and the rare flat ( <90 mm) abalone (   H. walallensis ) on protected and fished rocky reefs in California. Abalone abundance surveys were conducted inside 24 × 30 m plots on three protected reefs with red sea urchins present and three fished reefs where red sea urchins were removed by commercial or experimental fishing. Significantly more juvenile abalone were found in 1996 and 1997 on protected reefs with urchins present than on fished reefs ( χ   ² = 188, df = 1, p < 0.001 ). Juvenile red abalone abundance was not correlated with local adult red abalone abundance or habitat rugosity. One-third of the juveniles inside the MPAs were found under the urchins' spine canopy, as were a suite of unfished marine organisms. In the laboratory, juvenile abalone survived better (  χ   ² = 7.31, df = 1, p < 0.01) in crab predation experiments in which red sea urchins were available as shelter. Fishing red urchins reduced structural complexity, potentially decreasing microhabitat available for juvenile abalone. This example demonstrates how MPAs designed for one fished species may help other species, illustrating their usefulness for ecosystem-based fishery management and marine conservation.     

Assessing the current state of ecological connectivity in a large marine protected area system

The establishment of marine protected areas (MPAs) is a critical step in ensuring the continued persistence of marine biodiversity. Although the area protected in MPAs is growing, the movement of individuals (or larvae) among MPAs, termed connectivity, has only recently been included as an objective of many MPAs. As such, assessing connectivity is often neglected or oversimplified in the planning process. For promoting population persistence, it is important to ensure that protected areas in a system are functionally connected through dispersal or adult movement. We devised a multi-species model of larval dispersal for the Australian marine environment to evaluate how much local scale connectivity is protected in MPAs and determine whether the extensive system of MPAs truly functions as a network. We focused on non-migratory species with simplified larval behaviors (i.e., passive larval dispersal) (e.g., no explicit vertical migration) as an illustration. Of all the MPAs analyzed (approximately 2.7 million km²), outside the Great Barrier Reef and Ningaloo Reef, <50% of MPAs (46-80% of total MPA area depending on the species considered) were functionally connected. Our results suggest that Australia's MPA system cannot be referred to as a single network, but rather a collection of numerous smaller networks delineated by natural breaks in the connectivity of reef habitat. Depending on the dispersal capacity of the taxa of interest, there may be between 25 and 47 individual ecological networks distributed across the Australian marine environment. The need to first assess the underlying natural connectivity of a study system prior to implementing new MPAs represents a key research priority for strategically enlarging MPA networks. Our findings highlight the benefits of integrating multi-species connectivity into conservation planning to identify opportunities to better incorporate connectivity into the design of MPA systems and thus to increase their capacity to support long-term, sustainable biodiversity outcomes.     

Marine habitat mapping in support of Marine Protected Area management in a subarctic fjord: Gilbert Bay, Labrador, Canada

This paper presents an approach that allows production of benthic substrate and habitat maps in fjord environments. This approach is used to support the management of the Gilbert Bay Marine Protected Area (MPA) in southeastern Labrador, Atlantic Canada. Multibeam sonar-derived bathymetry, seabed slope, and acoustic reflectance (backscatter) were combined using supervised classification methods and GIS with ground-truthed benthic sampling in order to derive maps of the substrates and main benthic habitats. Six acoustically distinct substrate types were identified in the fjord, and three additional substrate types without a unique acoustic signature were recognized. Ordination by multidimensional scaling and analysis of similarity generalized these to four acoustically distinct habitat types. Greatest within-habitat (alpha) diversity was found in the coralline-algae encrusted gravel habitat. Greatest between-habitat (beta) diversity was found in the management Zones 1 and 2, which have the highest level of protection. The study confirmed that the zoning plan for the MPA, which was designed to protect spawning and juvenile fish habitat for a local genetically distinct population of Atlantic cod, afforded highest levels of protection to areas with highest habitat diversity.     

Use of Habitats as Surrogates of Biodiversity for Efficient Coral Reef Conservation Planning in Pacific Ocean Islands

Abstract: Marine protected areas (MPAs) have been highlighted as a means toward effective conservation of coral reefs. New strategies are required to more effectively select MPA locations and increase the pace of their implementation. Many criteria exist to design MPA networks, but generally, it is recommended that networks conserve a diversity of species selected for, among other attributes, their representativeness, rarity, or endemicity. Because knowledge of species’ spatial distribution remains scarce, efficient surrogates are urgently needed. We used five different levels of habitat maps and six spatial scales of analysis to identify under which circumstances habitat data used to design MPA networks for Wallis Island provided better representation of species than random choice alone. Protected‐area site selections were derived from a rarity–complementarity algorithm. Habitat surrogacy was tested for commercial fish species, all fish species, commercially harvested invertebrates, corals, and algae species. Efficiency of habitat surrogacy varied by species group, type of habitat map, and spatial scale of analysis. Maps with the highest habitat thematic complexity provided better surrogates than simpler maps and were more robust to changes in spatial scales. Surrogates were most efficient for commercial fishes, corals, and algae but not for commercial invertebrates. Conversely, other measurements of species‐habitat associations, such as richness congruence and composition similarities provided weak results. We provide, in part, a habitat‐mapping methodology for designation of MPAs for Pacific Ocean islands that are characterized by habitat zonations similar to Wallis. Given the increasing availability and affordability of space‐borne imagery to map habitats, our approach could appreciably facilitate and improve current approaches to coral reef conservation and enhance MPA implementation.     

Impact of marine protected areas on temporal stability of fish species diversity

Preserving biodiversity over time is a pressing challenge for conservation science. A key goal of marine protected areas (MPAs) is to maintain stability in species composition, via reduced turnover, to support ecosystem function. Yet, this stability is rarely measured directly under different levels of protection. Rather, evaluations of MPA efficacy generally consist of static measures of abundance, species richness, and biomass, and rare measures of turnover are limited to short-term studies involving pairwise (beta diversity) comparisons. Zeta diversity is a recently developed metric of turnover that allows for measurement of compositional similarity across multiple assemblages and thus provides more comprehensive estimates of turnover. We evaluated the effectiveness of MPAs at preserving fish zeta diversity across a network of marine reserves over 10 years in Batemans Marine Park, Australia. Snorkel transect surveys were conducted across multiple replicated and spatially interspersed sites to record fish species occurrence through time. Protection provided by MPAs conferred greater stability in fish species turnover. Marine protected areas had significantly shallower decline in zeta diversity compared with partially protected and unprotected areas. The retention of harvested species was four to six times greater in MPAs compared with partially protected and unprotected areas, and the stabilizing effects of protection were observable within 4 years of park implementation. Conversely, partial protection offered little to no improvement in stability, compared with unprotected areas. These findings support the efficacy of MPAs for preserving temporal fish diversity stability. The implementation of MPAs helps stabilize fish diversity and may, therefore, support biodiversity resilience under ongoing environmental change.     

Coral Reef Habitats as Surrogates of Species, Ecological Functions, and Ecosystem Services

Abstract: Habitat maps are often the core spatially consistent data set on which marine reserve networks are designed, but their efficacy as surrogates for species richness and applicability to other conservation measures is poorly understood. Combining an analysis of field survey data, literature review, and expert assessment by a multidisciplinary working group, we examined the degree to which Caribbean coastal habitats provide useful planning information on 4 conservation measures: species richness, the ecological functions of fish species, ecosystem processes, and ecosystem services. Approximately one‐quarter to one‐third of benthic invertebrate species and fish species (disaggregated by life phase; hereafter fish species) occurred in a single habitat, and Montastraea‐dominated forereefs consistently had the highest richness of all species, processes, and services. All 11 habitats were needed to represent all 277 fish species in the seascape, although reducing the conservation target to 95% of species approximately halved the number of habitats required to ensure representation. Species accumulation indices (SAIs) were used to compare the efficacy of surrogates and revealed that fish species were a more appropriate surrogate of benthic species (SAI = 71%) than benthic species were for fishes (SAI = 42%). Species of reef fishes were also distributed more widely across the seascape than invertebrates and therefore their use as a surrogate simultaneously included mangroves, sea grass, and coral reef habitats. Functional classes of fishes served as effective surrogates of fish and benthic species which, given their ease to survey, makes them a particularly useful measure for conservation planning. Ecosystem processes and services exhibited great redundancy among habitats and were ineffective as surrogates of species. Therefore, processes and services in this case were generally unsuitable for a complementarity‐based approach to reserve design. In contrast, the representation of species or functional classes ensured inclusion of all processes and services in the reserve network.     

The application of predicted habitat models to investigate the spatial ecology of demersal fish assemblages

Benthic habitats are known to influence the abundance and richness of demersal fish assemblages; however, little is known about how habitat structure and composition influences these distributions at very fine scales. We examined how the benthic environment structures marine fish assemblages using high-resolution bathymetry and accurate predicted benthic habitat maps. Areas characterised by a mosaic of habitat patches supported the highest richness of demersal fishes. A total of 37.4% of the variation in the distribution of the fish assemblage was attributed to 6 significant variables. Depth explained 23.0% of the variation, with the boulders explaining 12.6% and relief 1.4%. The remaining measures (seawhips, light/exposure and solid reef) provided a small (<1.0%) but significant contribution. Identifying components of the benthic environment important in structuring fish assemblages and understanding how they influence the spatial distribution of marine fishes is imperative for better management of demersal fish populations.     

Lag Effects in the Impacts of Mass Coral Bleaching on Coral Reef Fish, Fisheries, and Ecosystems

Abstract:  Recent episodes of coral bleaching have led to wide-scale loss of reef corals and raised concerns over the effectiveness of existing conservation and management efforts. The 1998 bleaching event was most severe in the western Indian Ocean, where coral declined by up to 90% in some locations. Using fisheries-independent data, we assessed the long-term impacts of this event on fishery target species in the Seychelles, the overall size structure of the fish assemblage, and the effectiveness of two marine protected areas (MPAs) in protecting fish communities. The biomass of fished species above the size retained in fish traps changed little between 1994 and 2005, indicating no current effect on fishery yields. Biomass remained higher in MPAs, indicating they were effective in protecting fish stocks. Nevertheless, the size structure of the fish communities, as described with size-spectra analysis, changed in both fished areas and MPAs, with a decline in smaller fish (<30 cm) and an increase in larger fish (>45 cm). We believe this represents a time-lag response to a reduction in reef structural complexity brought about because fishes are being lost through natural mortality and fishing, and are not being replaced by juveniles. This effect is expected to be greater in terms of fisheries productivity and, because congruent patterns are observed for herbivores, suggests that MPAs do not offer coral reefs long-term resilience to bleaching events. Corallivores and planktivores declined strikingly in abundance, particularly in MPAs, and this decline was associated with a similar pattern of decline in their preferred corals. We suggest that climate-mediated disturbances, such as coral bleaching, be at the fore of conservation planning for coral reefs.     

Effectiveness of Marine Protected Areas in the Philippines for Biodiversity Conservation

Abstract: Quantifying the extent to which existing reserves meet conservation objectives and identifying gaps in coverage are vital to developing systematic protected‐area networks. Despite widespread recognition of the Philippines as a global priority for marine conservation, limited work has been undertaken to evaluate the conservation effectiveness of existing marine protected areas (MPAs). Targets for MPA coverage in the Philippines have been specified in the 1998 Fisheries Code legislation, which calls for 15% of coastal municipal waters (within 15 km of the coastline) to be protected within no‐take MPAs, and the Philippine Marine Sanctuary Strategy (2004), which aims to protect 10% of coral reef area in no‐take MPAs by 2020. We used a newly compiled database of nearly 1000 MPAs to measure progress toward these targets. We evaluated conservation effectiveness of MPAs in two ways. First, we determined the degree to which marine bioregions and conservation priority areas are represented within existing MPAs. Second, we assessed the size and spacing patterns of reserves in terms of best‐practice recommendations. We found that the current extent and distribution of MPAs does not adequately represent biodiversity. At present just 0.5% of municipal waters and 2.7–3.4% of coral reef area in the Philippines are protected in no‐take MPAs. Moreover, 85% of no‐take area is in just two sites; 90% of MPAs are <1 km². Nevertheless, distances between existing MPAs should ensure larval connectivity between them, providing opportunities to develop regional‐scale MPA networks. Despite the considerable success of community‐based approaches to MPA implementation in the Philippines, this strategy will not be sufficient to meet conservation targets, even under a best‐case scenario for future MPA establishment. We recommend that implementation of community‐based MPAs be supplemented by designation of additional large no‐take areas specifically located to address conservation targets.     

Viability Analysis of Reef Fish Populations Based on Limited Demographic Information

Abstract:  Marine protected areas (MPAs) that allow some degree of artisanal fishing have been proposed to control the overexploitation of marine resources while allowing extraction by local communities. Nevertheless, the management of MPAs is often impaired by the absence of data on the status of their resources. We devised a method to estimate population growth rates with the type of data that are usually available for reef fishes. We used 7 years of spatially explicit abundance data on the leopard grouper ( Mycteroperca rosacea ) in an MPA in the Gulf of California, Mexico, to construct a matrix population model that incorporated the effects of El Niño/La Niña Southern Oscillation on population dynamics. An environmental model that estimated different demographic estimates for El Niño and La Niña periods performed better than a single-environment model, and a single-habitat model performed better than a model that considered different depths as different habitats. Our results suggest that the population of the leopard grouper off the main island of the MPA is not viable under present conditions. Although the impact of fishing on leopard grouper populations in the MPA has not yet been established, fishing should be closed as a precautionary measure at this island if a priority of the MPA is to ensure the sustainability of its fish populations.     

Protection from fishing alters the species composition of fish assemblages in a temperate-tropical transition zone

Closure of areas to fishing is expected to result in an increase in the abundance of targeted species; however, changes to populations of species not targeted by fishermen will depend upon their role in the ecosystem and their relationship with targeted species. The effects of protection on targeted and non-targeted reef fish species at the Houtman Abrolhos Islands, Western Australia were studied using baited remote underwater stereo–video cameras. Video images were collected from shallow (8–12 m) and deep (22–26 m) reef sites inside a Marine Protected Area (MPA) at each of three island groups and from three replicate fished locations at each of these groups that span a temperate-tropical transition area. The MPAs were established in 1994 and vary in size from 13.72 km² at the Pelsaert group in the south to 22.29 km² at the Easter group to 27.44 km² at the Wallabi group in the north. The relative abundances of 137 fish species from 42 families were recorded. Large differences in fish assemblage structure existed between MPA and fished locations, and also between shallow and deep regions. Targeted fish species Plectropomus leopardus, Lethrinus miniatus, Lethrinus nebulosus, Pagrus auratus and Glaucosoma hebraicum were more abundant inside MPAs than in areas open to fishing. Their abundance inside MPAs was between 1.13 and 8 times greater than their abundance at fished locations. For non-targeted fish species many were more abundant in areas open to fishing, e.g. Coris auricularis, Thalassoma lutescens, Thalassoma lunare, Dascyllus trimaculatus, however others were conversely more abundant inside MPAs, e.g. Gymnothorax spp, Kyphosus sydneyanus, Scarus microhinos, Chromis westaustralis, Chaetodon spp. This study demonstrates that the removal of abundant targeted species from an ecosystem by fishing can indirectly impact non-fished species and alter the trophic structure of fish assemblages. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Temperate marine reserves enhance targeted but not untargeted fishes in multiple no-take MPAs.

Although many papers report the effects of no-take marine protected areas (MPAs or reserves), scientifically rigorous empirical studies are rare, particularly for temperate reef fishes. We evaluated the responses of fish populations to protection from fishing in reserves by comparing densities and sizes inside and outside of five no-take reserves in southern California, USA. Our results are robust because we compared responses across multiple rocky-reef reserves in two different years and controlled for possible site differences by (a) ensuring that habitat characteristics were the same inside and outside reserves, and (b) sampling species that are not targeted, which would not be expected to have a direct response to fishing. We compared fish density and size and calculated biomass and egg production across all five sites. Fishes targeted by recreational and/or commercial fisheries consistently exhibited increases in mean density (150%), size (30%), biomass (440%), and egg production (730%) inside reserves. Reserve effects were greatest for legal-sized targeted fishes: significantly greater densities were found exclusively inside reserves for targeted species (580%), the largest size classes existed only inside reserves, and mean biomass was 1000% higher. These responses were unlikely to have been caused by habitat differences because there were no significant differences in habitat characteristics between reserve and control locations. Densities of non-targeted species did not differ between reserve and non-reserve locations, further supporting the conclusions that differences in targeted species between reserve and control locations were due to harvesting rather than site-specific effects. Although MPAs cannot replace traditional fisheries management, the concentration of increased biomass and egg production is a unique MPA benefit that serves both reserves and fisheries. Scientifically rigorous studies that include multiple reserves, such as this study, are needed to inform management and policy decisions.