Ecological success of no-take marine protected areas: Using population dynamics theory to inform a global meta-analysis.

Adaptively managing marine protected areas (MPAs) requires accurately assessing whether established MPAs are achieving their goals of protecting and conserving biomass, especially for harvested populations. Ecological MPA assessments commonly compare inside of the MPA to a reference point outside of and/or before implementation (i.e., calculating "response ratios"). Yet, MPAs are not simple ecological experiments; by design, protected populations interact with those outside, and population dynamic responses can be nonlinear. This complicates assessment interpretations. Here, we used a two-patch population model to explore how MPA response ratios (outside-inside, before-after, and before-after-control-impact [BACI]) for fished populations behave under different conditions, like whether the population is receiving a sustainable larval supply or if it is declining despite protection from harvest. We then conducted a Bayesian evaluation of MPA effects on fish and invertebrate populations based on data collected from 82 published studies on 264 no-take MPAs worldwide, using the results of an earlier global meta-analysis as priors. We considered the effects of calculating different summary metrics on these results, drawing on the theoretical insights from our population model as a comparative framework. We demonstrate that not all response ratio comparison types provide the same information: For example, outside-inside and BACI comparisons can fail to detect population decline within MPAs, whereas before-after comparisons likely detect that pattern. Considering these limitations, we nonetheless found that MPAs globally are producing positive outcomes, with on average greater biomass, density, and organism size within their boundaries than reference sites. However, only a small portion of studies (18 of 82) provided the temporal data necessary to determine that protection, on average, has led to increased abundance of populations within MPAs over time. These findings demonstrate the importance of considering the underlying system dynamics when assessing MPA effects. Assuming that large outside-inside or BACI response ratios always reflect large and net positive conservation effects may lead to misleading conclusions, we recommend that: (1) when assessing specific MPA effects, empirical findings be considered alongside theoretical knowledge relevant to that MPA system, and (2) management should respond to the local conditions and outcomes, rather than a blanket expectation for positive MPA effects.

No-take marine protected areas enhance the benefits of kelp-forest restoration for fish but not fisheries.

Kelp habitat restoration is gaining traction as a management action to support recovery in areas affected by severe disturbances, thereby ensuring the sustainability of ecosystem services. Knowing when and where to restore is a major question. Using a single-species population model, we consider how restoring inside marine protected areas (MPAs) might benefit coastal fish populations and fisheries. We found that MPAs can greatly enhance the population benefits of restoration but at a small cost to fishery yields. Generally, restoring inside MPAs had a better overall gains-loss outcome, especially if the system is under high fishing pressure or severe habitat loss. However, restoring outside became preferable when predatory fish indirectly benefit kelp habitats. In either case, successful restoration actions may be difficult to detect in time-series data due to complex transient dynamics. We provide context for setting management goals and social expectations for the ecosystem service implications of restoration in MPAs.

Recruitment variability and sampling design interact to influence the detectability of protected area effects.

Correctly identifying the effects of a human impact on a system is a persistent challenge in ecology, driven partly by the variable nature of natural systems. This is particularly true in many marine fishery species, which frequently experience large temporal fluctuations in recruitment that produce interannual variations in populations. This variability complicates efforts to maintain stocks at management targets or detect the effects of rebuilding efforts. We address this challenge in the context of no-take marine reserves by exploring how variable larval recruitment could interact with the timing of reserve establishment and choice of sampling design to affect population dynamics and the detectability of reserve effects. To predict population changes in the years following a no-take reserve implementation, we first tested for periodicity in larval recruitment in an important U.S. Pacific coast recreational fishery species (kelp bass, Paralabrax clathratus) and then included that pattern in a population model. We also used this model to determine the detectability of population increases under alternative sampling approaches and minimum age sampled. Kelp bass larval recruitment in the Channel Islands, California, peaked every about six (major) and about two (minor) years. Our model showed that establishing a reserve during a peak or trough enhanced or delayed, respectively, the post-reserve population increases. However, establishing a reserve during a recruitment peak could obscure a failing reserve, that is, a reserve that is unable to secure longer-term metapopulation persistence. Recruitment peaks and troughs also interacted with sampling design to affect the detectability of reserve effects. Designs that compared inside-outside were the most robust to variable recruitment, but failed to capture whether the reserve has improved metapopulation growth. Designs that included a time element (e.g., before-after) are more suited to assessing reserve effectiveness, but were sensitive to recruitment variation and detectability can change year-to-year. Notably, detectability did not always increase monotonically with reserve age; the optimal time for detectability depended on the minimum age of organisms sampled and was greatest when the cohort of a major recruitment peak first appeared in the sampling. We encourage managers to account for variable recruitment when planning monitoring and assessment programs.

Marine reserves stabilize fish populations and fisheries yields in disturbed coral reef systems.

Marine reserve networks are increasingly implemented to conserve biodiversity and enhance the persistence and resilience of exploited species and ecosystems. However, the efficacy of marine reserve networks in frequently disturbed systems, such as coral reefs, has rarely been evaluated. Here we analyze a well-mixed larval pool model and a spatially explicit model based on a well-documented coral trout (Plectropomus spp.) metapopulation in the Great Barrier Reef Marine Park, Australia, to determine the effects of marine reserve coverage and placement (in relation to larval connectivity and disturbance heterogeneity) on the temporal stability of fisheries yields and population biomass in environmentally disturbed systems. We show that marine reserves can contribute to stabilizing fishery yield while increasing metapopulation persistence, irrespective of whether reserves enhance or diminish average fishery yields. However, reserve placement and the level of larval connectivity among subpopulations were important factors affecting the stability and sustainability of fisheries and fish metapopulations. Protecting a mix of disturbed and non-disturbed reefs, rather than focusing on the least-disturbed habitats, was the most consistently beneficial approach across a range of dispersal and reserve coverage scenarios. Placing reserves only in non-disturbed areas was the most beneficial for biomass enhancement, but had variable results for fisheries and could potentially destabilize yields in systems with well-mixed larval or those that are moderately fished. We also found that focusing protection on highly disturbed areas could actually increase variability in yields and biomass, especially when degraded reef reserves were distant and poorly connected to the meta-population. Our findings have implications for the design and implementation of reserve networks in the presence of stochastic, patchy environmental disturbances.

Fishery consequences of marine reserves: short-term pain for longer-term gain.

Marine reserves are often established in areas that support fisheries. Larval export from reserves is argued to help compensate for the loss of fishable habitat; however, previous modeling studies have focused on long-term equilibrium outcomes. We examined the transient consequences of reserve establishment for fished metapopulations, considering both a well-mixed larval pool and a spatially explicit model based on a coral trout (Plectropomus spp.) metapopulation. When fishing pressure was reallocated relative to the area protected, yields decreased initially, then recovered, and ultimately exceeded pre-reserve levels. However, recovery time was on the order of several years to decades. If fishing pressure intensified to maintain pre-reserve yields, reserves were sometimes unable to support the increased mortality and the metapopulation collapsed. This was more likely when reserves were small, or located peripherally within the metapopulation. Overall, reserves can achieve positive conservation and fishery benefits, but fisheries management complementary to reserve implementation is essential.

Planning Marine Reserve Networks for Both Feature Representation and Demographic Persistence Using Connectivity Patterns.

Marine reserve networks must ensure the representation of important conservation features, and also guarantee the persistence of key populations. For many species, designing reserve networks is complicated by the absence or limited availability of spatial and life-history data. This is particularly true for data on larval dispersal, which has only recently become available. However, systematic conservation planning methods currently incorporate demographic processes through unsatisfactory surrogates. There are therefore two key challenges to designing marine reserve networks that achieve feature representation and demographic persistence constraints. First, constructing a method that efficiently incorporates persistence as well as complementary feature representation. Second, incorporating persistence using a mechanistic description of population viability, rather than a proxy such as size or distance. Here we construct a novel systematic conservation planning method that addresses both challenges, and parameterise it to design a hypothetical marine reserve network for fringing coral reefs in the Keppel Islands, Great Barrier Reef, Australia. For this application, we describe how demographic persistence goals can be constructed for an important reef fish species in the region, the bar-cheeked trout (Plectropomus maculatus). We compare reserve networks that are optimally designed for either feature representation or demographic persistence, with a reserve network that achieves both goals simultaneously. As well as being practically applicable, our analyses also provide general insights into marine reserve planning for both representation and demographic persistence. First, persistence constraints for dispersive organisms are likely to be much harder to achieve than representation targets, due to their greater complexity. Second, persistence and representation constraints pull the reserve network design process in divergent directions, making it difficult to efficiently achieve both constraints. Although our method can be readily applied to the data-rich Keppel Islands case study, we finally consider the factors that limit the method's utility in information-poor contexts common in marine conservation.

Synergistic Effects of Marine Reserves and Harvest Controls on the Abundance and Catch Dynamics of a Coral Reef Fishery.

Marine no-take reserves, where fishing and other extractive activities are prohibited, have well-established conservation benefits [1], yet their impacts on fisheries remains contentious [2-4]. For fishery species, reserves are often implemented alongside more conventional harvest strategies, including catch and size limits [2, 5]. However, catch and fish abundances observed post-intervention are often attributed to reserves, without explicitly estimating the potential contribution of concurrent management interventions [2, 3, 6-9]. Here we test a metapopulation model against observed fishery [10] and population [11] data for an important coral reef fishery (coral trout; Plectropomus spp.) in Australia's Great Barrier Reef Marine Park (GBRMP) to evaluate how the combined increase in reserve area [12] and reduction in fishing effort [13, 14] in 2004 influenced changes in fish stocks and the commercial fishery. We found that declines in catch, increases in catch rates, and increases in biomass since 2004 were substantially attributable to the integration of direct effort controls with the rezoning, rather than the rezoning alone. The combined management approach was estimated to have been more productive for fish and fisheries than if the rezoning had occurred alone and comparable to what would have been obtained with effort controls alone. Sensitivity analyses indicate that the direct effort controls prevented initial decreases in catch per unit effort that would have otherwise occurred with the rezoning. Our findings demonstrate that by concurrently restructuring the fishery, the conservation benefits of reserves were enhanced and the fishery cost of rezoning the reserve network was socialized, mitigating negative impacts on individual fishers.