Expanding marine protected areas to include degraded coral reefs.

Marine protected areas (MPAs) are a commonly applied solution to coral reef degradation, yet coral reefs continue to decline worldwide. We argue that expanding the range of MPAs to include degraded reefs (DR-MPA) could help reverse this trend. This approach requires new ecological criteria for MPA design, siting, and management. Rather than focusing solely on preserving healthy reefs, our approach focuses on the potential for biodiversity recovery and renewal of ecosystem services. The new criteria would help identify sites with the highest potential for recovery and the greatest resistance to future threats (e.g., increased temperature and acidification) and sites that contribute to MPA connectivity. The DR-MPA approach is a compliment rather than a substitute for traditional MPA design approaches. We believe that the DR-MPA approach can enhance the natural, or restoration-assisted, recovery of DRs and their ecosystem services; increase total reef area available for protection; promote more resilient and better-connected MPA networks; and improve conditions for human communities dependent on MPA ecosystem services.

From principles to practice: a spatial approach to systematic conservation planning in the deep sea.

Increases in the demand and price for industrial metals, combined with advances in technological capabilities have now made deep-sea mining more feasible and economically viable. In order to balance economic interests with the conservation of abyssal plain ecosystems, it is becoming increasingly important to develop a systematic approach to spatial management and zoning of the deep sea. Here, we describe an expert-driven systematic conservation planning process applied to inform science-based recommendations to the International Seabed Authority for a system of deep-sea marine protected areas (MPAs) to safeguard biodiversity and ecosystem function in an abyssal Pacific region targeted for nodule mining (e.g. the Clarion-Clipperton fracture zone, CCZ). Our use of geospatial analysis and expert opinion in forming the recommendations allowed us to stratify the proposed network by biophysical gradients, maximize the number of biologically unique seamounts within each subregion, and minimize socioeconomic impacts. The resulting proposal for an MPA network (nine replicate 400 × 400 km MPAs) covers 24% (1 440 000 km(2)) of the total CCZ planning region and serves as example of swift and pre-emptive conservation planning across an unprecedented area in the deep sea. As pressure from resource extraction increases in the future, the scientific guiding principles outlined in this research can serve as a basis for collaborative international approaches to ocean management.

Detecting larval export from marine reserves.

Marine reserve theory suggests that where large, productive populations are protected within no-take marine reserves, fished areas outside reserves will benefit through the spillover of larvae produced in the reserves. However, empirical evidence for larval export has been sparse. Here we use a simple idealized coastline model to estimate the expected magnitude and spatial scale of larval export from no-take marine reserves across a range of reserve sizes and larval dispersal scales. Results suggest that, given the magnitude of increased production typically found in marine reserves, benefits from larval export are nearly always large enough to offset increased mortality outside marine reserves due to displaced fishing effort. However, the proportional increase in recruitment at sites outside reserves is typically small, particularly for species with long-distance (on the order of hundreds of kilometers) larval dispersal distances, making it very difficult to detect in field studies. Enhanced recruitment due to export may be detected by sampling several sites at an appropriate range of distances from reserves or at sites downcurrent of reserves in systems with directional dispersal. A review of existing empirical evidence confirms the model's suggestion that detecting export may be difficult without an exceptionally large differential in production, short-distance larval dispersal relative to reserve size, directional dispersal, or a sampling scheme that encompasses a broad range of distances from the reserves.

The stochastic nature of larval connectivity among nearshore marine populations.

Many nearshore fish and invertebrate populations are overexploited even when apparently coherent management structures are in place. One potential cause of mismanagement may be a poor understanding and accounting of stochasticity, particularly for stock recruitment. Many of the fishes and invertebrates that comprise nearshore fisheries are relatively sedentary as adults but have an obligate larval pelagic stage that is dispersed by ocean currents. Here, we demonstrate that larval connectivity is inherently an intermittent and heterogeneous process on annual time scales. This stochasticity arises from the advection of pelagic larvae by chaotic coastal circulations. This result departs from typical assumptions where larvae simply diffuse from one site to another or where complex connectivity patterns are created by transport within spatially complicated environments. We derive a statistical model for the expected variability in larval settlement patterns and demonstrate how larval connectivity varies as a function of different biological and physical processes. The stochastic nature of larval connectivity creates an unavoidable uncertainty in the assessment of fish recruitment and the resulting forecasts of sustainable yields.

Fish in offshore kelp forests affect recruitment to intertidal barnacle populations.

Kelp forests along the coast of central California harbor juvenile rockfish that prey on the larvae of invertebrates from the rocky intertidal zone. This predation reduces recruitment to barnacle populations to 1/50 of the level in the absence of fish. The dynamics of the intertidal community are thus strongly coupled to the dynamics of the offshore kelp community.

Local scale genetic structure in coral populations: a signature of selection.

Coastal marine reserves in general, and coral reef reserves in particular, are typically composed of scattered patches separated by uninhabited areas. Due to the sessile mode of life of adult corals, larval connectivity is often the only agent of gene flow between reef localities. In this study we examined the connectivity between populations of the common scleratinian coral Stylophora pistillata at the northern tip of the Gulf of Aqaba (Red Sea), using the rDNA ITS (internal transcribed spacer) as a molecular marker. Sequence comparisons among recruits indicated very similar, equally-diverse, assemblages of recruits in both the northern (highly affected by anthropogenic disturbances) and southern (less affected) study sites, implying a high larval connectivity or common sources of larval supply. By contrast, sequence diversity observed among adults declined sharply from southern to northern sites, accompanied by genetic differentiation of the respective populations. Based on Fu's Fs-test of selective neutrality, it may be suggested that various post-settlement selective regimes, presumably more intense in the northern sites, provide a reasonable explanation for the observed patterns of genetic diversity. The suggested hypothesis is supported by the sharper decline in sequence diversity found between recruits and adults in the northern sites. This study exemplifies the necessity to consider local selective factors, in addition to larval connectivity, when managing marine reserves.

Marine spatial planning makes room for offshore aquaculture in crowded coastal waters.

Marine spatial planning (MSP) seeks to reduce conflicts and environmental impacts, and promote sustainable use of marine ecosystems. Existing MSP approaches have successfully determined how to achieve target levels of ocean area for particular uses while minimizing costs and impacts, but they do not provide a framework that derives analytical solutions in order to co-ordinate siting of multiple uses while balancing the effects of planning on each sector in the system. We develop such a framework for guiding offshore aquaculture (bivalve, finfish, and kelp farming) development in relation to existing sectors and environmental concerns (wild-capture fisheries, viewshed quality, benthic pollution, and disease spread) in California, USA. We identify > 250,000 MSP solutions that generate significant seafood supply and billions of dollars in revenue with minimal impacts (often < 1%) on existing sectors and the environment. We filter solutions to identify candidate locations for high-value, low-impact aquaculture development. Finally, we confirm the expectation of substantial value of our framework over conventional planning focused on maximizing individual objectives.

A comparative study of asymmetric migration events across a marine biogeographic boundary.

In many nonclonal, benthic marine species, geographic distribution is mediated by the dispersal of their larvae. The dispersal and recruitment of marine larvae may be limited by temperature gradients that can affect mortality or by ocean currents that can directly affect the movements of pelagic larvae. We focus on Point Conception, a well-known biogeographic boundary between the Californian and Oregonian biogeographic provinces, to investigate whether ocean currents affect patterns of gene flow in intertidal marine invertebrates. The predominance of pelagically dispersing species with northern range limits at Point Conception suggests that ocean currents can affect species distributions by erecting barriers to the dispersal of planktonic larvae. In this paper, we investigate whether the predominantly southward currents have left a recognizable genetic signature in species with pelagically dispersing larvae whose ranges span Point Conception. We use patterns of genetic diversity and a new method for inferring cladistic migration events to test the hypothesis that southward currents increase southward gene flow for species with pelagically dispersing larvae. We collected mitochondrial DNA (mtDNA) sequence data for the barnacles Balanus glandula and Chthamalus fissus and also reanalyzed a previously published mtDNA dataset (Strongylocentrotus purpuratus, Edmands et al. 1996). For all three species, our cladistic approach identified an excess of southward migration events across Point Conception. In data from a fourth species with nondispersing larvae (Nucella emarginata, Marko 1998), our method suggests that ocean currents have not played a role in generating genetic structure.

'Heads I win, tails you lose': testing directional alternative hypotheses in ecological and evolutionary research.

Whenever experiments make a priori predictions about the direction of change in some parameter, one-tailed test statistics offer a potentially large gain in power over the corresponding two-tailed test. This gain is rarely used in ecology and evolution because of (1) the belief that one-tailed procedures are unavailable for most statistical tests and (2) an inherent dilemma in one-tailed tests: how do we handle large parameter changes in the unanticipated direction? The first problem is a misconception, whereas the second is easily resolved by recognizing that one- and two-tailed tests are simply extremes in a continuum of testing options.

One-way analysis of variance with unequal variances.

We have designed a statistical test that eliminates the assumption of equal group variances from one-way analysis of variance. This test is preferable to the standard technique of trial-and-error transformation and can be shown to be an extension of the Behrens-Fisher T test to the case of three or more means. We suggest that this procedure be used in most applications where the one-way analysis of variance has traditionally been applied to biological data.

Extending nondirectional heterogeneity tests to evaluate simply ordered alternative hypotheses.

Biologists frequently use nondirectional heterogeneity tests when comparing three or more populations because a suitable directional test is unavailable or is not practical to implement. Here we describe a test, the ordered heterogeneity test, that permits testing against simply ordered alternative hypotheses in the context of almost any nondirectional test. The test has a wide range of parametric and nonparametric applications. Graphs are developed for calculating exact P values.

Coastal oceanography sets the pace of rocky intertidal community dynamics.

The structure of ecological communities reflects a tension among forces that alter populations. Marine ecologists previously emphasized control by locally operating forces (predation, competition, and disturbance), but newer studies suggest that inputs from large-scale oceanographically modulated subsidies (nutrients, particulates, and propagules) can strongly influence community structure and dynamics. On New Zealand rocky shores, the magnitude of such subsidies differs profoundly between contrasting oceanographic regimes. Community structure, and particularly the pace of community dynamics, differ dramatically between intermittent upwelling regimes compared with relatively persistent down-welling regimes. We suggest that subsidy rates are a key determinant of the intensity of species interactions, and thus of structure in marine systems, and perhaps also nonmarine communities.