Effects of fishing pressure and trophic group on abundance and spillover across boundaries of a no-take zone

We investigated factors influencing fish abundance and emigration across the boundaries of a no-take zone (NTZ), and its adjacent fished zones, within the Nabq Managed Resource Protected Area, South Sinai, Egyptian Red Sea. Underwater visual census of eight families of reef fish was undertaken at three depths across the whole NTZ (1.2 km), and for a similar distance into the adjacent fished zones. Because most fishing occurs in shallow water, the effect of the NTZ changed with depth. On reef flat transects (1 m depth), seven families showed significantly higher abundance in the NTZ, whereas at 10 m depth only two herbivorous families, Acanthuridae (surgeonfishes) and Siganidae (rabbitfishes) showed a significant difference, both being more abundant in the fished zones, an effect most likely due to reduced competition or predation. To look for evidence of emigration (spillover) of fish from the NTZ, data were also tested for evidence of gradients in abundance extending from within the NTZ to within the fished zones. In shallow water six families showed significant declines in abundance moving away from the centre of the NTZ, whereas at 10 m, the only significant gradients were for acanthurids and siganids, both becoming more abundant moving further into the fished zones. Comparisons of estimated fish length between areas also showed significant differences. The results indicate that at Nabq, while fishing reduces the abundance of most families, especially predators, abundance of some families and species, especially herbivores, may increase under light fishing pressure. Thus, spillover may occur to an extent and in a direction depending on trophic group and fishing intensity.     

Impacts of lost fishing gear on coral reef sessile invertebrates in the Florida Keys National Marine Sanctuary

The Florida Keys coral reef ecosystem supports multimillion-dollar commercial and recreational fisheries. The ecological effects caused by fishing gear that is lost when cut or broken after snagging on the bottom is a growing concern to managers and scientists. Few data exist, however, to assess the impacts of lost fishing gear to benthic organisms and habitat structure. In this study, 63 offshore coral reef and hard-bottom sites were surveyed during 2001 to quantify the impacts of lost fishing gear to coral reef sessile invertebrates. Lost hook-and-line fishing gear accounted for 87% of all debris (N=298 incidences) encountered and was responsible for 84% of the 321 documented impacts to sponges and benthic cnidarians, predominantly consisting of tissue abrasion causing partial individual or colony mortality. Branching gorgonians (Octocorallia) were the most frequently affected (56%), followed by milleporid hydrocorals (19%) and sponges (13%). Factors affecting the impacts of lost fishing gear include sessile invertebrate density, the density of lost fishing gear, and gear length. While lost hook-and-line fishing gear is ubiquitous in the Florida Keys, less than 0.2% of the available milleporid hydrocorals, stony corals, and gorgonians in the habitats studied are adversely affected in terms of colony abrasions and partial mortality.     

Effectiveness of five small Philippines’ coral reef reserves for fish populations depends on site-specific factors, particularly enforcement history

Community-managed, no-take marine reserves are increasingly promoted as a simple, precautionary measure to conserve biodiversity and sustain coral reef fisheries. However, we need to demonstrate the effects of such reserves to those affected by the loss of potential fishing grounds and the wider scientific community. We surveyed changes in fish communities in five small marine reserves in the central Philippines and three distant Control sites over seven years. We conducted underwater visual censuses of 53 fish families within the reserve (Inside), with a kilometre of the boundary (Outside) and at Control sites. We found significant differences between fish communities Inside and Outside the reserve only at the two sites with strictest compliance with fishing prohibition, while there were significant differences to distant Control sites in all cases. The strongest responses to reserve protection were found in predatory fishes (groupers and breams) and in butterflyfish. Other abundant fish families showed weak effects of protection. For all taxa analysed, we found significant effects of reserve Site and Site × Treatment interactions. The detection of fish responses to reserves is complicated by potential spillover effects, site-specific factors, particularly compliance, and the difficulty of identifying appropriate control areas.     

Effects of fishing protection on the genetic structure of fish populations

Marine reserves have been identified as an important tool in the management of fishery resources and their number is increasing rapidly, most of them being on islands. However, knowledge on the real effect of protection from fishing on the genetic structure of populations, the spatial scales involved, or the suitability of islands as reserves in terms of connectivity, is scarce. This paper analyses the effects of fishery protection on the genetic structure of populations of Diplodus sargus, a target species, in protected and non-protected areas of the western Mediterranean. Populations studied showed high genetic variability at spatial scales from 10¹ to 10³ km. Protected areas have significantly higher allelic richness. The lower levels of heterozygosis and higher heterozygote deficit showed by islands compared with coastal areas makes clear the importance of considering the connectivity processes when designing a MPA.     

Community-based conservation results in the recovery of reef fish spawning aggregations in the Coral Triangle

In the Coral Triangle community-based marine protected areas (MPAs) are being established at a prolific rate. Their establishment can benefit both fisheries and biodiversity, and they provide both a socially and economically acceptable means of managing coral reefs in developing nations. However, because such MPAs are typically small (usually <0.5 km²), they will rarely provide protection to large mobile fishes. An exception to this limitation may exist when community-based MPAs are established to protect small sites where vital processes occur, such as fish spawning aggregations (FSAs). To test the effectiveness of small (0.1–0.2 km²) MPAs for protecting FSAs, we monitored three FSA sites where brown-marbled grouper (Epinephelus fuscoguttatus), camouflage grouper (Epinephelus polyphekadion) and squaretail coralgrouper (Plectropomus areolatus) aggregate to spawn. Sites were monitored during peak reproductive periods (several days prior to each new moon) between January 2005 and November 2009. All three sites are located in New Ireland Province, Papua New Guinea, and had been exploited for decades, but in 2004 two sites were protected by the establishment of community-based MPAs. The third site continued to be exploited. Over the monitoring period densities of E. fuscoguttatus and E. polyphekadion increased at both MPAs, but not at the site that remained open to fishing. At one MPA the densities of E. polyphekadion increased tenfold. Our findings demonstrate that community-based MPAs that are appropriately designed and adequately enforced can lead to the recovery of populations of vulnerable species that aggregate to spawn.     

Effects of fishing on sex-changing Caribbean parrotfishes

We studied parrotfish (Scaridae) assemblages on coral reefs in relation to fishing pressure around six Caribbean islands. Fishing intensity ranged from virtually none in Bonaire, and increased through Saba, Puerto Rico, St Lucia and Dominica to extremely high levels in Jamaica. In St Lucia we also compared parrotfish assemblages between fishing grounds and fully protected marine reserves, from 1995, 6 months prior to establishment, to 2001. Within each country we performed replicate counts of the number and size of all parrotfish species within, or passing through our counting area. From these data we calculated biomass for seven species. Biomass of the two largest species, Sparisoma viride and Scarus vetula, was greatest in islands with low fishing pressure (P<0.001). By contrast, smaller species constituted an increasing proportion of the total parrotfish assemblage as fishing pressure increased (P<0.001 in all cases). Parrotfish are protogynous hermaphrodites with two distinct colour phases. The initial phase is predominantly female, and the terminal phase exclusive to sexually mature males. The average size of all species except Sc. vetula tended to decrease with increasing fishing pressure. Furthermore, percentages of fish that were terminal phase males showed order of magnitude declines with increasing fishing pressure for Sp. viride and Sc. vetula. Terminal males of these species were absent from counts in Jamaica and virtually absent from Dominica suggesting that persistence of these populations may depend on recruitment from distant sources. Following reserve implementation in St Lucia, all species, except uncommon Sp. chrysopterum, increased in mean biomass (P<0.001 in all cases). In 6 years the total biomass for all species combined increased to become nearly four times as high in reserves and almost twice as high in fishing grounds [P<0.001 (year effect); P<0.001 (protection effect); P<0.001 (year×protection)], and mean size of five species increased significantly in both reserves and fishing grounds.     

Assessing the effects of marine protected area (MPA) on a reef fish assemblage in a northwestern Mediterranean marine reserve: Identifying community-based indicators

Marine protected areas (MPAs) are increasingly envisaged as a tool to manage coastal ecosystems and fisheries. Assessment of their performance with respect to management objectives is therefore important. A number of MPAs provided conservation benefits for fished species. Observed benefits do not apply to all species at all times, and responses to protection are also highly variable among fish taxa. Among the many empirical studies on marine reserves, only a few designs considered ‘before and after data’ and spatial variation. In this paper, we are interested in assessing the effect of a no-take reserve on the reef fish assemblage in a northwestern Mediterranean example. Data were obtained from a three-year survey using underwater visual censuses (UVC), before and after MPA establishment. Permutational multivariate analysis of variance (PERMANOVA) and multivariate regression trees (MRT) were used to evaluate the effects of reserve protection on the reef fish assemblage, while accounting for habitat. Modelled biological responses were abundances and diversity indices calculated at different levels of the assemblage. Significant effects were found for many of these metrics. In addition to PERMANOVA, univariate models provided more insight into the magnitude and direction of effects. The most sensitive metrics were related to large species and species targeted by fishing. These results may be used to choose the metrics that are more suitable as community-based indicators of MPA impact in the perspective of monitoring programs.     

Reefs of an uninhabited Caribbean island: fishes, benthic habitat, and opportunities to discern reef fishery impact

Navassa Island is a tiny, (5 km²) uninhabited US protectorate located between Jamaica and Haiti. It is part of the Caribbean Islands National Wildlife Refuge, under the jurisdiction of the US Fish and Wildlife Service. We conducted a quantitative assessment of Navassa's coral reef fishes and benthic habitat, in order to assist with the development of conservation plan for the island. The shallow reefs of Navassa (<23m) have high live coral cover (range 20-26.1%), high degree of architectural complexity (rugosity index range 1.4-1.9), and moderate abundance of the keystone grazing urchin, Diadema antillarum, at all sites (mean 2.9±0.9 per 30 m²). Despite its remoteness, an unregulated, artisanal fishery (primarily using traps and hook and line) carried out by Haitians is the primary mode of human impact on Navassa reefs. Even so, reef fish communities exhibit high density (range 97-140 fish per 60 m²) and retain representation by large snapper, grouper and herbivores, which are mostly lacking in nearby Caribbean locations with high fishing pressure. Thus, Navassa reefs appear to be trophically intact with fish populations relatively “unexploited,” presenting a conservation challenge and a research opportunity. The regulation and conservation of the fishery will be difficult, due to the international nature of the situation. However, given the apparently small impact that artisanal fisheries have yet had on its reef communities, Navassa presents a possibly unique opportunity to study the ecological functioning of a relatively trophically intact Caribbean reef, and represents a strong imperative for conservation, monitoring, and research.     

A quantitative method for evaluating the importance of marine areas for conservation of birds

Objective criteria are needed for ranking marine sites when examining candidate areas for protection measures. We suggest a Marine Classification Criterion (MCC) which allows the application of the widely used Ramsar 1% criterion for wetlands for seabirds with clustered distribution in offshore habitats. The maximum size of an area considered to be internationally important has not been defined by the Ramsar Convention. Terrestrial and coastal sites generally have obvious hydrological or physical boundaries, whereas such boundaries are less obvious at sea. The smallest unit which would pass the demands set by the MCC is 1% of the bio-geographic population of a particular species concentrated in an area (site) supporting a density exceeding a value equivalent of four times the average density of the species in the investigated regional sea. The effect of choosing smaller or larger reference densities is tested. The results indicate that the chosen threshold density is a suitable requirement for the inclusion of the most important areas for seabird species with at least 25% of their bio-geographic population occurring in the studied regions of the North Sea and the Baltic Sea. The test cases indicate that provided the MCC is based on geo-statistical analyses of un-biased survey data the boundaries of areas holding large concentrations of seabirds can be estimated with confidence. The MCC could be used to identify concentrations of seabirds and other marine animals of conservation priority and to rank marine areas by their cumulative importance to different species.     

Alleviating impacts of anthropogenic activities by traditional conservation measures: can a small reef reserve be sustainedly managed?

Zoning schemes that are becoming an important management tool in large marine reserves, are difficult to implement in small reef areas. At the 3.4 km long reef of Eilat (Red Sea), a small (ca. 350 m of coastline) enclosure strategy has been enforced since 1992, while the remaining reef was left open to intense human activities. Here we have investigated for 2.5 y three populations of the branching coral Stylophora pistillata (3605 colonies) in a locality within the enclosed area (site NR) and in two areas open to the public, by tossing random quadrats at the shallow lagoonar zone (0.5-1.5 m depth). In the two open sites we found significantly higher levels of colony breakage (14-34% vs. 4-9% in the enclosed site), lower partial mortality levels of colonies (in the first 1.5 y; 7-9% vs. 23-30% at NR), higher recruitment (up to 3.0 vs. up to 0.9 colonies/m 2), 50% reduction in coral life span (10 vs. 20 y) and an estimated extinction period of 9-10 y for new cohorts as compared to >20 y in the enclosed site. Average colony size and maximal colony size were about half in the open sites. Live coverage fluctuated widely in all sites but was 3 times higher in the enclosed area (1.0-3.0% vs. 0.3-1.1%). Log-transformed size frequency distributions revealed, at the open sites, a shift from small towards medium-size classes and at the enclosed site, a shift from larger to medium size classes. We conclude that the enclosure of a limited core zone, although improved some ecological parameters, was not sufficient to compensate for stress imposed by anthropogenic activities. It is suggested to employ active restoration approaches, such as the “gardening concept”, as supplementary management tools.     

Patterns of protection and threats along productivity gradients in Canada

Productivity is an important driver of broad-scale diversity gradients and community composition. Surprisingly, it is rarely used as a biodiversity proxy in protected area network assessments. In this research, we evaluate if biases exist in the locations of Canada’s protected areas with respect to productivity and assess the distribution of anthropogenic threats to protection along productivity and topographic axes. Productivity was expressed as the annual minimum and integrated fraction of absorbed photosynthetically active radiation (fPAR) and related seasonality (defined as variation in fPAR over a year), as acquired by remote sensing. Results indicate that, overall, protected areas are slightly biased to lower productivities (higher seasonalities), but are relatively unbiased along elevation. Sites at high elevations, high seasonalities, or low productivities are extremely well represented, but they account for a very small proportion of Canada’s area overall and under protection. However, the productivity characteristics of protected areas are heterogeneous across reserve characteristics and space; biases are greater and more variable when considering individual International Union for Conservation of Nature (IUCN) categories, protected area size classes, or ecozones.The spatial and environmental correlates of anthropogenic pressures (proximity of roads and settlements in the greater park ecosystem) to protected areas in the forested ecozones were evaluated with partial linear regression models. Threats were strongly spatially structured. Reserve characteristics (productivity, elevation, size, IUCN category) shared about half of this spatial structure, explaining ∼25% of the variation in threat distributions. Small and productive protected areas tended to occur closer to human populations and roads, respectively, and are thus expected to face greater threats to biodiversity. High-elevation protected areas generally occurred farther from both roads and settlements.Systematic patterns between productivity, reserve size and management goals, and anthropogenic disturbances suggest that the most productive, biodiverse areas may not yet be sufficiently protected. These analyses highlight considerations for the management of existing parks and the expansion of Canada’s protected area network.     

Fishers’ perception of a 35-year old exclusive Fisheries Management Zone

Fishers’ attitudes and perceptions are critical for the success of fisheries protection areas with their associated biota, and a failure to understand fishers’ behaviour may undermine the success of such fisheries management measures. In this study, we examine fishers’ perception of a long-established exclusive fisheries zone around Malta and to investigate if the perceptions depend on fishers’ demographic, economic, social characteristics and fishing activity of the fishers. A questionnaire survey was undertaken to evaluate the demographic characteristics, economic situation (costs and revenue) and fishers’ activity and behaviour, together with their perception of the Fisheries Management Zone (FMZ). A total of 241 interview responses were analysed which was a response rate of 60%. The perception of most fishers was that the establishment of the FMZ has had an overall negative impact on their fishing activity and that the zone is not important for the protection of local fish stocks. When asked about the beneficial effect of the zone for fishers, most fishers from all backgrounds said that the zone does not benefit commercial fishers, but benefits mainly recreational fishers. The most evident differences in the perceptions and attitudes were between the full-time, part-time and recreational fishers. Fishers that have been fishing for more than 35 years and fishers from the main fishing village also had different attitudes from other fishers towards the FMZ. The results of this study suggest that the proportion of individual income derived from fishing was the strongest factor that influenced attitudinal differences, with home port and fishing experience having less important effects. The main differences in attitude among fishers were related to the protection and conservation effects of the zone, enhancement of resources and conflicts among user groups. The heterogeneity among fishers’ attitudes revealed by the present study has important implications for the implementation of spatial closures. Some sectors of stakeholders may require additional incentives to accept restrictions on access if spatial management is to achieve its intended objectives.     

A fuzzy logic expert system to estimate intrinsic extinction vulnerabilities of marine fishes to fishing

Fishing has become a major conservation threat to marine fishes. Effective conservation of threatened species requires timely identification of vulnerable species. However, evaluation of extinction risk using conventional methods is difficult for the majority of fish species because the population data normally required by such methods are unavailable. This paper presents a fuzzy expert system that integrates life history and ecological characteristics of marine fishes to estimate their intrinsic vulnerability to fishing. We extract heuristic rules (expressed in IF-THEN clauses) from published literature describing known relationships between biological characteristics and vulnerability. Input and output variables are defined by fuzzy sets which deal explicitly with the uncertainty associated with qualitative knowledge. Conclusions from different lines of evidence are combined through fuzzy inference and defuzzification processes. Our fuzzy system provides vulnerability estimates that correlate with observed declines more closely than previous methods, and has advantages in flexibility of input data requirements, in the explicit representation of uncertainty, and in the ease of incorporating new knowledge. This fuzzy expert system can be used as a decision support tool in fishery management and marine conservation planning.     

The importance of spatial scale for the conservation of tidal flat macrobenthos: An example from New South Wales, Australia

Planners of Marine Protected Areas (MPAs) commonly use maps of habitat types when choosing areas to conserve. This assumes that habitats are homogeneous, and therefore, that any area of habitat will represent the full spectrum of ecological diversity within that habitat. Here, we report that macrobenthic assemblages in tidal flat habitats were spatially heterogeneous in terms of beta diversity (taxonomic turnover), abundance, taxonomic richness and Shannon-Wiener H′ Diversity. Importantly, the patterns of heterogeneity were scale dependent for the three spatial scales we examined; plots (20 m), sites (100s of m) and estuaries (<30 km). The three estuaries in the study were compositionally similar as they shared the same dominant taxa, although one estuary had significantly more taxa and a higher abundance of macrobenthos. Assemblages within tidal flats differed at scales of 100s of m for all ecological measures. Most notably, beta diversity was highest at this scale. Assemblages were relatively more homogeneous at the 20-m scale. These findings highlight the value of examining more than one ecological measure and estimating magnitude of effects across a variety of scales. This work presents two important considerations for MPAs. First, although tidal flats in different estuaries are compositionally similar for dominant taxa, rarer taxa and high heterogeneity in abundance should influence the choice and number of tidal flats in MPAs. Second, strong compositional heterogeneity within individual tidal flats implies that conservation of whole habitat, rather than sections of a tidal flat, is essential if this habitat type is to be used to represent taxonomic diversity.     

Inter-annual variability in the home range of breeding turtles: Implications for current and future conservation management

We assessed home range size for breeding loggerhead turtles (Caretta caretta) near the limit of the species range at the Greek island of Zakynthos in the Mediterranean. Thirteen adult females and seven adult males were tracked using GPS units (loggers and transmitters) during May and June of 2006, 2007 and 2008. Kernel analysis indicated that core home range sizes (50% estimator; range: 2.9-19.7 km²) for both males and females were restricted to a 7.5 km tract of coastline. 15% of GPS locations fell outside of the national park protection zones, while within the protected breeding area 88% of GPS locations occurred in zones of minimal protection. Female home ranges were 64% larger in 2008 than in 2006 and 2007, indicating that several years monitoring may be required for the most effective designation of marine protected areas (MPAs). Ten of the tracked females departed the core breeding area on 15 occasions for periods of 1-15 days travelling distances of 10-100 km, although none nested at alternative breeding sites. The inter-annual variability of breeding area home range size and likelihood of incidence of forays appeared be correlated with barometric pressure. The movement responses of loggerheads to environmental conditions implicates an ability to switch nesting areas over small scales in response to climate change. However, such behaviour suggests the protection of existing core breeding sites may be inadequate, with policy makers being required to consider the protection of broader areas to encompass potential changes in the habitat needs of this species.     

Shortcuts for marine conservation planning: The effectiveness of socioeconomic data surrogates

Marine protected area (MPA) networks designed without consideration of the interests of local communities are likely to fail. However, in many regions where conservation action is needed most urgently, socioeconomic data are not available at spatial scales relevant to conservation planning. In the Philippines, the primary stakeholders relevant to conservation efforts in coastal waters are small-scale fishers. Unlike commercial fisheries, no logbook data are kept to record fishers’ spatial effort and usage patterns. We investigated the effects of including different surrogates for small-scale fishing effort in the systematic design of an MPA network for Siquijor Province. We compared a reserve selection scenario in which socioeconomic data were not considered with four different surrogates for fishing effort and with empirical data on the spatial distribution of fishing effort collected through interviews. We assumed that minimising opportunity costs to fishers would increase the likelihood that they would support and comply with MPA implementation, resulting in more effective conservation. Surrogates modelled on the number of fishers or boats in each community consistently outperformed those based on population census data. However, none of the surrogates we tested were able to accurately predict fine-scale resource use patterns. Whilst socioeconomic surrogates may be able to assist conservation planners to identify regional-scale opportunities where conservation objectives may be met more easily, they cannot act as a shortcut for comprehensive consultation with communities, which will be required to identify actual sites for MPA implementation.     

Testing the utility of abiotic surrogates for marine habitat mapping at scales relevant to management

Habitat mapping at the scale at which marine protected areas are designed and managed is essential for assessment of, and design for, representation. Most habitat mapping studies rely solely or in part on abiotic surrogates for patterns of biodiversity. The utility of abiotic variables in predicting biological distributions at the local scale (10 s of km) was tested in a remote video survey of macrobenthos in Moreton Bay, Australia. Habitat classifications of the same set of 41 sites based on 6 abiotic variables and abundances of 89 taxa and bioturbation indicators were compared using correlation, regression and ordination analyses. The concepts of false homogeneity (abiotically similar but biologically distinct) and false heterogeneity (abiotically distinct but biologically similar) were defined to describe types of errors associated with using abiotic surrogates to construct habitat maps, and quantified using two separate methods. Overall, the best prediction by abiotic surrogates explained less than 30% of the pattern of biological similarity. Errors of false homogeneity were between 20% and 62%, depending on the methods of estimation. Predictive capability of abiotic surrogates at the taxon level was poor, with only 6% of taxon/surrogate correlations significant. Abiotic variables did not discriminate sufficiently between different soft bottom communities to be a reliable basis for mapping. These results have implications for the widespread use of abiotic surrogates in marine habitat mapping to plan for, or assess, representation in marine protected areas. Little confidence can be placed in marine habitat classifications based solely or largely on abiotic surrogates without calibration by rigorous biological surveys at the appropriate scale. Therefore, it is questionable whether marine protected areas designed on this basis can have measurable benefits for conservation.     

Entanglement of monofilament fishing lines and coral death

This study examined the extent of damage by monofilament fishing lines on the cauliflower coral (Pocillopora meandrina) colonies at a popular cast fishing site in Oahu, Hawaii in March 1998. Sixty-five percent of coral colonies had fishing lines on their surface, and 80% of colonies were either entirely or partially dead. The percentage of entirely and partially dead colonies was positively correlated with the percentage of colonies entangled with fishing lines. Percent dead coral surface area was larger for colonies with fishing lines than colonies without fishing lines. In colonies with fishing lines, percent dead coral surface area was positively correlated with percent area with fishing lines. The high incidence of coral colonies with fishing lines and the three sets of significant results indicated destructive effects of monofilament fishing lines to P. meandrina corals. There is thus a clear case that reefs should be protected not only from the effects of excessive removal of fishes, but also from the effects of unsuccessful fishing that results in fishing-line entanglement on corals.     

Consequences of adult and juvenile movement for marine protected areas

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