Measuring the long-term success of small-scale marine protected areas in a Philippine reef fishery

Tropical coral reefs are subject to multiple pressures from both natural and anthropogenic sources. These pressures have caused widespread declines in reef health, resulting in the increased use of spatial management tools such as marine protected areas (MPAs). MPAs have proven generally effective if well designed and enforced, but there are limited long-term studies investigating how the presence of small-scale MPAs affects fish populations and reef communities. Using a 12-year time series, we found that small-scale (10–50 ha) community-managed MPAs along the Danajon Bank of the Philippines preserved average fish biomass within their boundaries over time relative to surrounding fished reefs. Unprotected areas are, however, showing significant long-term biomass decline. MPAs were also found to preserve more key trophic groups and larger-bodied commercially targeted reef fish families. Fish biomass of piscivore, scavenger and invertivore trophic groups inside individual MPAs is, however, still declining at a similar rate as outside. Surprisingly, long-term benthic cover and growth form composition were not significantly affected overall by MPA presence, despite the sporadic use of highly destructive dynamite fishing in this region. Coral cover has remained historically low (21–28%) throughout the study, following widespread bleaching mortality. While management tempered overall abundance declines, we found that irrespective of MPA presence, there was a generalised decline of both large- and small-bodied fish size groups across the study region, most steeply within the 20–30 cm length fish, and a shift towards proportionally higher abundances of small (5–10 cm) fish. This indicates a combination of over-exploitation, inadequate MPA size and coverage for larger fish, and the lingering effects of the 1998 bleaching event. Generalised shifts in body size and trophic structure reported here could lead to future reductions in fishery productivity and stability and will be further exacerbated unless broader fishery regulations and enforcement is instated.

     

Rapid Effects of Marine Reserves via Larval Dispersal

Marine reserves have been advocated worldwide as conservation and fishery management tools. It is argued that they can protect ecosystems and also benefit fisheries via density-dependent spillover of adults and enhanced larval dispersal into fishing areas. However, while evidence has shown that marine reserves can meet conservation targets, their effects on fisheries are less understood. In particular, the basic question of if and over what temporal and spatial scales reserves can benefit fished populations via larval dispersal remains unanswered. We tested predictions of a larval transport model for a marine reserve network in the Gulf of California, Mexico, via field oceanography and repeated density counts of recently settled juvenile commercial mollusks before and after reserve establishment. We show that local retention of larvae within a reserve network can take place with enhanced, but spatially-explicit, recruitment to local fisheries. Enhancement occurred rapidly (2 yrs), with up to a three-fold increase in density of juveniles found in fished areas at the downstream edge of the reserve network, but other fishing areas within the network were unaffected. These findings were consistent with our model predictions. Our findings underscore the potential benefits of protecting larval sources and show that enhancement in recruitment can be manifested rapidly. However, benefits can be markedly variable within a local seascape. Hence, effects of marine reserve networks, positive or negative, may be overlooked when only focusing on overall responses and not considering finer spatially-explicit responses within a reserve network and its adjacent fishing grounds. Our results therefore call for future research on marine reserves that addresses this variability in order to help frame appropriate scenarios for the spatial management scales of interest.     

When Is Spillover from Marine Reserves Likely to Benefit Fisheries?

The net movement of individuals from marine reserves (also known as no-take marine protected areas) to the remaining fishing grounds is known as spillover and is frequently used to promote reserves to fishers on the grounds that it will benefit fisheries. Here we consider how mismanaged a fishery must be before spillover from a reserve is able to provide a net benefit for a fishery. For our model fishery, density of the species being harvested becomes higher in the reserve than in the fished area but the reduction in the density and yield of the fished area was such that the net effect of the closure was negative, except when the fishery was mismanaged. The extent to which effort had to exceed traditional management targets before reserves led to a spillover benefit varied with rates of growth and movement of the model species. In general, for well-managed fisheries, the loss of yield from the use of reserves was less for species with greater movement and slower growth. The spillover benefit became more pronounced with increasing mis-management of the stocks remaining available to the fishery. This model-based result is consistent with the literature of field-based research where a spillover benefit from reserves has only been detected when the fishery is highly depleted, often where traditional fisheries management controls are absent. We conclude that reserves in jurisdictions with well-managed fisheries are unlikely to provide a net spillover benefit.     

Empirical Evidence for Species-Specific Export of Fish Na?veté from a No-Take Marine Protected Area in a Coastal Recreational Hook and Line Fishery

No-take marine protected areas (MPAs) are assumed to enhance fisheries catch via the “spillover” effect, where biomass is exported to adjacent exploited areas. Recent studies in spearfishing fisheries suggest that the spillover of gear-naïve individuals from protected to unprotected sites increases catch rates outside the boundaries of MPAs. Whether this is a widespread phenomenon that also holds for other gear types and species is unknown. In this study, we tested if the distance to a Mediterranean MPA predicted the degree of vulnerability to hook and line in four small-bodied coastal fish species. With the assistance of underwater video recording, we investigated the interaction effect of the distance to the boundary of an MPA and species type relative to the latency time to ingest a natural bait, which was considered as a surrogate of fish naïveté or vulnerability to fishing. Vulnerability to angling increased (i.e., latency time decreased) within and near the boundary of an MPA for an intrinsically highly catchable species (Serranus scriba), while it remained constant for an intrinsically uncatchable control species (Chromis chromis). While all of the individuals of S. scriba observed within the MPA and surrounding areas were in essence captured by angling gear, only one fifth of individuals in the far locations were captured. This supports the potential for the spillover of gear-naïve and consequently more vulnerable fish from no-take MPAs. Two other species initially characterized as intermediately catchable (Coris julis and Diplodus annularis) also had a shorter latency time in the vicinity of an MPA, but for these two cases the trend was not statistically significant. Overall, our results suggest that an MPA-induced naïveté effect may not be universal and may be confined to only intrinsically highly catchable fish species. This fact emphasizes the importance of considering the behavioural dimension when predicting the outcomes of MPAs, otherwise the effective contribution may be smaller than predicted for certain highly catchable species such as S. scriba.     

Spillover of fish naïveté from marine reserves

Spillover of adult fish biomass is an expected benefit from no‐take marine reserves to adjacent fisheries. Here, we show fisher‐naïve behaviour in reef fishes also spills over from marine reserves, potentially increasing access to fishery benefits by making fishes more susceptible to spearguns. The distance at which two targeted families of fishes began to flee a potential fisher [flight initiation distance (FID)] was lower inside reserves than in fished areas, and this reduction extended outside reserve boundaries. Reduced FID persisted further outside reserves than increases in fish biomass. This finding could help increase stakeholder support for marine reserves and improve current models of spillover by informing estimates for spatial changes in catchability. Behavioural changes of fish could help explain differences between underwater visual census and catch data in quantifying the spatial extent of spillover from marine reserves, and should be considered in the management of adjacent fisheries.     

Assessing the Contribution of Marine Protected Areas to the Trophic Functioning of Ecosystems: A Model for the Banc d’Arguin and the Mauritanian Shelf

Most modelling studies addressed the effectiveness of marine protected areas (MPA) for fisheries sustainability through single species approach. Only a few models analysed the potential benefits of MPAs at the ecosystem level, estimating the potential export of fish biomass from the reserve or analysing the trophic relationships between organisms inside and outside the MPA. Here, we propose to use food web models to assess the contribution of a MPA to the trophic functioning of a larger ecosystem. This approach is applied to the Banc d’Arguin National Park, a large MPA located on the Mauritanian shelf. The ecosystem was modeled using Ecopath with Ecosim, a model that accounts for fisheries, food web structure, and some aspects of the spatial distribution of species, for the period 1991–2006. Gaps in knowledge and uncertainty were taken into account by building three different models. Results showed that the Banc d’Arguin contributes about 9 to 13% to the total consumption, is supporting about 23% of the total production and 18% of the total catch of the Mauritanian shelf ecosystem, and up to 50% for coastal fish. Of the 29 exploited groups, 15 depend on the Banc for more than 30% of their direct or indirect consumptions. Between 1991 and 2006, the fishing pressure increased leading to a decrease in biomass and the catch of high trophic levels, confirming their overall overexploitation. Ecosim simulations showed that adding a new fleet in the Banc d’Arguin would have large impacts on the species with a high reliance on the Banc for food, resulting in a 23% decrease in the current outside MPA catches. We conclude on the usefulness of food web models to assess MPAs contribution to larger ecosystem functioning.     

Assessing Dispersal Patterns of Fish Propagules from an Effective Mediterranean Marine Protected Area

Successfully enforced marine protected areas (MPAs) have been widely demonstrated to allow, within their boundaries, the recovery of exploited species and beyond their boundaries, the spillover of juvenile and adult fish. Little evidence is available about the so-called ‘recruitment subsidy’, the augmented production of propagules (i.e. eggs and larvae) due to the increased abundance of large-sized spawners hosted within effective MPAs. Once emitted, propagules can be locally retained and/or exported elsewhere. Patterns of propagule retention and/or export from MPAs have been little investigated, especially in the Mediterranean. This study investigated the potential for propagule production and retention/export from a Mediterranean MPA (Torre Guaceto, SW Adriatic Sea) using the white sea bream, Diplodus sargus sargus, as a model species. A multidisciplinary approach was used combining 1) spatial distribution patterns of individuals (post-settlers and adults) assessed through visual census within Torre Guaceto MPA and in northern and southern unprotected areas, 2) Lagrangian simulations of dispersal based on an oceanographic model of the region and data on early life-history traits of the species (spawning date, pelagic larval duration) and 3) a preliminary genetic study using microsatellite loci. Results show that the MPA hosts higher densities of larger-sized spawners than outside areas, potentially guaranteeing higher propagule production. Model simulations and field observation suggest that larval retention within and long-distance dispersal across MPA boundaries allow the replenishment of the MPA and of exploited populations up to 100 km down-current (southward) from the MPA. This pattern partially agrees with the high genetic homogeneity found in the entire study area (no differences in genetic composition and diversity indices), suggesting a high gene flow. By contributing to a better understanding of propagule dispersal patterns, these findings provide crucial information for the design of MPAs and MPA networks effective to replenish fish stocks and enhance fisheries in unprotected areas.     

Differential Response of Fish Assemblages to Coral Reef-Based Seaweed Farming

As the global demand for seaweed-derived products drives the expansion of seaweed farming onto shallow coral ecosystems, the effects of farms on fish assemblages remain largely unexplored. Shallow coral reefs provide food and shelter for highly diverse fish assemblages but are increasingly modified by anthropogenic activities. We hypothesized that the introduction of seaweed farms into degraded shallow coral reefs had potential to generate ecological benefits for fish by adding structural complexity and a possible food source. We conducted 210 transects at 14 locations, with sampling stratified across seaweed farms and sites adjacent to and distant from farms. At a seascape scale, locations were classified by their level of exposure to human disturbance. We compared sites where (1) marine protected areas (MPAs) were established, (2) neither MPAs nor blast fishing was present (hence “unprotected”), and (3) blast fishing occurred. We observed 80,186 fish representing 148 species from 38 families. The negative effects of seaweed farms on fish assemblages appeared stronger in the absence of blast fishing and were strongest when MPAs were present, likely reflecting the positive influence of the MPAs on fish within them. Species differentiating fish assemblages with respect to seaweed farming and disturbance were typically small but also included two key target species. The propensity for seaweed farms to increase fish diversity, abundance, and biomass is limited and may reduce MPA benefits. We suggest that careful consideration be given to the placement of seaweed farms relative to MPAs.     

Using marine reserves to estimate fishing mortality

The proportion of a fish stock that is killed by fishing activity is often calculated as the catch divided by the estimated stock biomass. However, stock biomass is notoriously difficult to estimate reliably, and moreover, the catch may be uncertain or misreported and does not include losses due to discarding. In all too many fisheries, these difficulties have lead to underestimates of total fishing mortality and the commercial demise of the fishery. No‐take marine reserves eliminate fishing mortality from within their boundaries and, for species that exhibit seasonal migratory behaviour, comparison of reserves with fished areas can provide direct estimates of the proportion killed by fishing. For an important exploited species in New Zealand, seasonal changes in density of sub‐legal fish at three marine reserves were similar in both reserve and adjacent non‐reserve areas. However, this result did not hold for legal‐size fish, and the difference in seasonal change between reserved and non‐reserved areas was used to obtain direct estimates of the total localized fishing mortality in the non‐reserve area over 6‐month periods. Estimates of the percentage of legal‐size fish killed by fishing ranged from 70 to 96%. These results demonstrate an unanticipated practical benefit from marine reserves that goes beyond their ecological role.     

Reef Fishes at All Trophic Levels Respond Positively to Effective Marine Protected Areas

Marine Protected Areas (MPAs) offer a unique opportunity to test the assumption that fishing pressure affects some trophic groups more than others. Removal of larger predators through fishing is often suggested to have positive flow-on effects for some lower trophic groups, in which case protection from fishing should result in suppression of lower trophic groups as predator populations recover. We tested this by assessing differences in the trophic structure of reef fish communities associated with 79 MPAs and open-access sites worldwide, using a standardised quantitative dataset on reef fish community structure. The biomass of all major trophic groups (higher carnivores, benthic carnivores, planktivores and herbivores) was significantly greater (by 40% - 200%) in effective no-take MPAs relative to fished open-access areas. This effect was most pronounced for individuals in large size classes, but with no size class of any trophic group showing signs of depressed biomass in MPAs, as predicted from higher predator abundance. Thus, greater biomass in effective MPAs implies that exploitation on shallow rocky and coral reefs negatively affects biomass of all fish trophic groups and size classes. These direct effects of fishing on trophic structure appear stronger than any top down effects on lower trophic levels that would be imposed by intact predator populations. We propose that exploitation affects fish assemblages at all trophic levels, and that local ecosystem function is generally modified by fishing.     

Estimating historical commercial rock lobster (Jasus edwardsii) catch inside Australian State territorial waters for marine protected area assessment: the binomial likelihood method

The rock lobster (Jasus edwardsii) fishery of South Australia is the State’s most valuable fisheries resource with an export value exceeding ~AU$100 million. The fishery operates primarily inshore (<60 m), and overlaps with a series of marine protected areas (MPAs) currently proposed for State territorial waters. As a result, the need to quantify the impact of proposed MPAs on commercial landings of rock lobster within territorial waters is an integral part of the MPA assessment process. Removing fishing effort displaced by MPAs prevents a corresponding increase in exploitation outside protected zones. We describe a binomial likelihood method that utilises historical commercial catch data to estimate catch totals of rock lobster inside South Australian State waters. Lobster catches per km² showed a high level of spatial variation with estimated historical lobster catch in State waters spanning approximately three orders of magnitude. The method identified key areas where high lobster catch (up to 500 kg/km²) overlapped with State waters. Binomial likelihood outputs have particular application to the estimation of net catch loss in situations where fishery buy-back or financial compensation are a considered option as part of the MPA implementation process.     

Spatial Heterogeneity in Fishing Creates de facto Refugia for Endangered Celtic Sea Elasmobranchs

The life history characteristics of some elasmobranchs make them particularly vulnerable to fishing mortality; about a third of all species are listed by the IUCN as Threatened or Near Threatened. Marine Protected Areas (MPAs) have been suggested as a tool for conservation of elasmobranchs, but they are likely to be effective only if such populations respond to fishing impacts at spatial-scales corresponding to MPA size. Using the example of the Celtic Sea, we modelled elasmobranch biomass (kg h⁻¹) in fisheries-independent survey hauls as a function of environmental variables and ‘local’ (within 20 km radius) fishing effort (h y⁻¹) recorded from Vessel Monitoring Systems data. Model selection using AIC suggested strongest support for linear mixed effects models in which the variables (i) fishing effort, (ii) geographic location and (iii) demersal fish assemblage had approximately equal importance in explaining elasmobranch biomass. In the eastern Celtic Sea, sampling sites that occurred in the lowest 10% of the observed fishing effort range recorded 10 species of elasmobranch including the critically endangered Dipturus spp. The most intensely fished 10% of sites had only three elasmobranch species, with two IUCN listed as Least Concern. Our results suggest that stable spatial heterogeneity in fishing effort creates de facto refugia for elasmobranchs in the Celtic Sea. However, changes in the present fisheries management regime could impair the refuge effect by changing fisher''s behaviour and displacing effort into these areas.     

Differential movement and movement bias models for marine protected areas

Marine protected areas (MPAs) are promoted as a tool to protect overfished stocks and increase fishery yields. Previous models suggested that adult mobility modified effects of MPAs by reducing densities of fish inside reserves, but increasing yields (i.e., increasing densities outside of MPAs). Empirical studies contradicted this prediction: as mobility increased, the relative density of fishes inside MPAs (relative to outside) increased or stayed constant. We hypothesized that this disparity between theoretical and empirical results was the result of differential movement of fish inside versus outside the MPA. We, therefore, developed a model with unequal and discontinuous diffusion, and analyzed its steady state and stability. We determined the abundance in the fishing grounds, the yield, the total abundance and the log ratio at steady-state and examined their response to adult mobility (while keeping the relative inequity in the diffusion constant). Abundance in the fishing grounds and yield increased, while total abundance and log-ratio decreased, as mobility increased. These results were all qualitatively consistent with the previous models assuming uniform diffusivity. Thus, the mismatch between empirical and theoretical results must result from other processes or other forms of differential movement. Therefore, we modified our original model by assuming that species located on the boundary of the MPA will preferentially move towards the MPA. This localized movement bias model gives rise to steady state profiles that can differ radically from the profiles in the unbiased model, especially when the bias is large. Moreover, for sufficiently large bias values, the monotonicity of the four measures with increased mobility is reversed, when compared with our original model. Thus, the movement bias model reconciles empirical data and theoretical results.     

Effects of reef-top gathering and fishing on invertebrate abundance across take and no-take zones

Gathering of molluscs on the reef-top, largely by women, is part of the traditional fishery practised by Bedouin in South Sinai, Egypt. The catch is dominated by Tridacna spp. and Octopus spp. Within the Nabq Managed Resource Protected Area, on the southern Gulf of Aqaba coast, a network of no-take zones (NTZs) was established (in 1995) to promote sustainable management of finfish stocks. Since this zoning also applied to exploitation of invertebrates, surveys of large species of molluscs and echinoderms across selected NTZs and adjacent fished areas were conducted (over 2000-2002) to assess any effects of gathering. Pooled data from all three years showed significantly higher abundances of Tridacna and Tectus dentatus in the NTZs, with greater abundances occurring at the reef edge zone. Size-frequency distributions revealed both a greater size range of Tridacna and a greater mean size of both Tridacna and Te. dentatus, within the NTZs, as compared to the fished areas. The size-frequency distribution of gleaned Tridacna, determined from discarded shells, also showed a smaller size range than did live Tridacna within the NTZs. By contrast, holothurians, which are not currently exploited by the local Bedouin, showed greater abundance in fished areas than in NTZs. Large diadematid sea urchins, along with the non-commercial strombs, Strombus gibberulus and Strombus fasciatus, were also significantly more abundant within fished areas, an effect presumed due to reduced predation pressure from fish as a result of fishing. Within the fished areas, there was a positive relationship between local abundance of Tridacna and of diadematids, thought to be due to high densities of the urchins acting as a deterrent against gathering. The establishment of NTZs has led to significant differences in invertebrate populations between take zones (TZ) and no-take zones. However, since there can be little or no spillover of adults of less mobile or sedentary invertebrates from NTZs to fished areas, any benefit to the fishery will depend largely on the greater larval production and export from protected populations.     

The effects of fishing on the diversity,biomass and trophic structure of Seychelles’ reef fish communities

A fishery independent underwater visual census technique was used to assess the effects of fishing on the diversity, biomass and trophic structure of the diurnally active non-cryptic reef-associated fish communities of the Seychelles. One hundred and thirty four species associated with three significantly different types of reef habitat were censused at one unfished ground and in six fishing grounds subject to different fishing intensity and the biomass of several species targeted by the fishery. The diversity of families containing target species (lutjanidae, lethrinidae) was significantly higher at unfished and lightly fished sites as was the total biomass of the fish community and the biomass of piscivorous, piscivorous/invertebrate feeding and herbivorous trophic groups. However, there was no indication that the biomass of non-target species increased in response to the removal of their predators by fishing. The findings of this study are significant for fishery managers because they suggest that the intensive differential cropping of top predators will not necessarily lead to increases in the biomass and productivity of their prey.     

Variation in reef fish and invertebrate communities with level of protection from fishing across the Eastern Tropical Pacific seascape

Aim To quantify general differences in reef community structure between well‐enforced and poorly enforced marine protected areas (MPAs) and fished sites across the Eastern Tropical Pacific (ETP) regional seascape Location The Pacific continental margin and oceanic islands of Costa Rica, Panama, Colombia and Ecuador, including World Heritage sites at Galapagos, Coiba, Cocos and Malpelo Methods Densities of reef fishes, mobile and sessile invertebrates, and macroalgae were quantified using underwater visual surveys at 136 ‘no‐take’ and 54 openly fished sites associated with seven large MPAs that encompassed a range of management strategies. Spatial variation in multivariate and univariate community metrics was related to three levels of fishing pressure (high‐protection MPAs, limited‐protection MPAs, fishing zones) for both continental and oceanic reefs. Results High‐protection MPAs possessed a much greater biomass of higher carnivorous fishes, lower densities of asteroids and Eucidaris spp. urchins, and higher coral cover than limited‐protection MPAs and fished zones. These results were generally consistent with the hypothesis that overfishing of predatory fishes within the ETP has led to increased densities of habitat‐modifying macroinvertebrates, which has contributed to regional declines in coral cover. Major differences in ecological patterns were also evident between continental and oceanic biogeographic provinces. Main conclusions Fishing down the food web, with associated trophic cascades, has occurred to a greater extent along the continental coast than off oceanic islands. Poorly enforced MPAs generate food webs more similar to those present in fished areas than in well‐protected MPAs.     

Long-term changes in deep-water fish populations in the northeast Atlantic: a deeper reaching effect of fisheries?

A severe scarcity of life history and population data for deep-water fishes is a major impediment to successful fisheries management. Long-term data for non-target species and those living deeper than the fishing grounds are particularly rare. We analysed a unique dataset of scientific trawls made from 1977 to 1989 and from 1997 to 2002, at depths from 800 to 4800 m. Over this time, overall fish abundance fell significantly at all depths from 800 to 2500 m, considerably deeper than the maximum depth of commercial fishing (approx. 1600 m). Changes in abundance were significantly larger in species whose ranges fell at least partly within fished depths and did not appear to be consistent with any natural factors such as changes in fluxes from the surface or the abundance of potential prey. If the observed decreases in abundance are due to fishing, then its effects now extend into the lower bathyal zone, resulting in declines in areas that have been previously thought to be unaffected. A possible mechanism is impacts on the shallow parts of the ranges of fish species, resulting in declines in abundance in the lower parts of their ranges. This unexpected phenomenon has important consequences for fisheries and marine reserve management, as this would indicate that the impacts of fisheries can be transmitted into deep offshore areas that are neither routinely monitored nor considered as part of the managed fishery areas.     

Larval export from marine reserves and the recruitment benefit for fish and fisheries

Marine reserves, areas closed to all forms of fishing, continue to be advocated and implemented to supplement fisheries and conserve populations. However, although the reproductive potential of important fishery species can dramatically increase inside reserves, the extent to which larval offspring are exported and the relative contribution of reserves to recruitment in fished and protected populations are unknown. Using genetic parentage analyses, we resolve patterns of larval dispersal for two species of exploited coral reef fish within a network of marine reserves on the Great Barrier Reef. In a 1,000 km(2) study area, populations resident in three reserves exported 83% (coral trout, Plectropomus maculatus) and 55% (stripey snapper, Lutjanus carponotatus) of assigned offspring to fished reefs, with the remainder having recruited to natal reserves or other reserves in the region. We estimate that reserves, which account for just 28% of the local reef area, produced approximately half of all juvenile recruitment to both reserve and fished reefs within 30 km. Our results provide compelling evidence that adequately protected reserve networks can make a significant contribution to the replenishment of populations on both reserve and fished reefs at a scale that benefits local stakeholders.     

Modeling the trophic effects of marine protected area zoning policies: A case study

Marine protected areas (MPAs) are increasingly being recognized as an alternative management tool for conserving marine resources and ecosystems. By integrating organism dispersal rates, ecosystem interactions and fishing effort dynamics, ECOSPACE, a spatially explicit ecosystem-based modeling tool, allowed us to compare the ecological consequences of alternative MPA zoning policies within the proposed Gwaii Haanas National Marine Conservation Area, located off the west coast of British Columbia, Canada. The desired effects of MPAs include higher fishery yields, the conservation of biodiversity, and/or the preservation of intact ecosystems. However, ECOSPACE predicts that when MPAs are small, species interactions and movements may make these objectives difficult to achieve. ECOSPACE suggests that the effects of MPAs are reduced at their boundaries where fishing effort is predicted to concentrate. Furthermore, top predators may become more abundant within MPAs, which could lead to a depression of their prey species and a subsequent increase of species at even lower trophic levels. Trophic cascade patterns and density gradients across boundaries are nontrivial departures from our simple expectations of how MPAs protect areas and will force us to reconsider what constitutes effective conservation. Our ECOSPACE model indicates that the establishment of multi-use buffer zones may help alleviate these realistic but worrisome ecological predictions. When coupled with an overall reduction in harvest pressure, ECOSPACE suggests that a MPA with a large core `no-take' zone and large buffer will result in the greatest increase in organism biomass. The use of marine zoning may be an effective management tactic to reduce social conflict and conserve marine ecosystems.     

Modelling spatial and temporal scales for spill-over and biomass exportation from MPAs and their potential for fisheries enhancement

Marine protected areas (MPAs) are considered as a tool for marine conservation and sustainable fishery resource management. Improvements in fishery yields should take place via the spill-over of individuals from the reserve. In general, it has been demonstrated that MPAs affect the density and biomass of the organisms within them, however, little evidence has been found in order to assess the exportation of individuals across their boundaries. In this study, a simple model involving population growth, harvest, and the diffusion coefficient for individuals was used to explore the effects of protection on populations inside the reserve and the spill-over of individuals to the fished area. The model showed that biological responses inside marine reserves appear to develop quickly, reaching mean levels within a short (1–5 year) time period. Mean population abundance is always higher inside the reserve and highlights the effectiveness of protection, particularly when there is strong fishing pressure outside the reserve. However, reserves smaller than 2000 m radius show significantly lower levels of abundance inside than larger sites. Large MPAs (i.e. about 2000 m in radius) offer nearly the maximum capacity for recovery (close to 100% of the system carrying capacity) and nearly the maximum flux of individuals per unit boundary length. Very large MPAs (i.e. larger than 6000 m in radius) could be a guaranteed means of providing resilience in order to prevent population crises, with the added advantage that the flux of individuals is slightly higher at larger distances from the boundary. However, in practice they provide no further advantage towards increasing the density of individuals or the exportation of biomass, and a network of smaller MPAs could be more beneficial, both from the point of view of conservation and of benefits to fisheries.