Potential of a no‐take marine reserve to protect home ranges of anadromous brown trout (Salmo trutta)

The extent to which no‐take marine reserves can benefit anadromous species requires examination. Here, we used acoustic telemetry to investigate the spatial behavior of anadromous brown trout (sea trout, Salmo trutta) in relation to a small marine reserve (~1.5 km²) located inside a fjord on the Norwegian Skagerrak coast. On average, sea trout spent 42.3 % (±5.0% SE) of their time in the fjord within the reserve, a proportion similar to the area of the reserve relative to that of the fjord. On average, sea trout tagged inside the reserve received the most protection, although the level of protection decreased marginally with increasing home range size. Furthermore, individuals tagged outside the reserve received more protection with increasing home range size, potentially opposing selection toward smaller home range sizes inflicted on fish residing within reserves, or through selective fishing methods like angling. Monthly sea trout home ranges in the marine environment were on average smaller than the reserve, with a mean of 0.430 (±0.0265 SE) km². Hence, the reserve is large enough to protect the full home range of some individuals residing in the reserve. Synthesis and applications: In general, the reserve protects sea trout to a varying degree depending on their individual behavior. These findings highlight evolutionary implications of spatial protection and can guide managers in the design of marine reserves and networks that preserve variation in target species' home range size and movement behavior.     

Effects of marine reserve characteristics on the protection of fish populations: a meta-analysis

Meta-analyses of published data for 19 marine reserves reveal that marine protected areas enhance species richness consistently, but their effect on fish abundance is more variable. Overall, there was a slight (11%) but significant increase in fish species number inside marine reserves, with all reserves sharing a common effect. There was a substantial but non-significant increase in overall fish abundance inside marine reserves compared to adjacent, non-reserve areas. When only species that are the target of fisheries were considered, fish abundance was significantly higher (by 28%) within reserve boundaries. Marine reserves vary significantly in the extent and direction of their response. This variability in relative abundance was not attributable to differences in survey methodology among studies, nor correlated with reserve characteristics such as reserve area, years since protection, latitude nor species diversity. The effectiveness of marine reserves in enhancing fish abundance may be largely related to the intensity of exploitation outside reserve boundaries and to the composition of the fish community within boundaries. It is recommended that studies of marine reserve effectiveness should routinely report fishing intensity, effectiveness of enforcement and habitat characteristics.     

Larval dispersal and movement patterns of coral reef fishes,and implications for marine reserve network design

Well‐designed and effectively managed networks of marine reserves can be effective tools for both fisheries management and biodiversity conservation. Connectivity, the demographic linking of local populations through the dispersal of individuals as larvae, juveniles or adults, is a key ecological factor to consider in marine reserve design, since it has important implications for the persistence of metapopulations and their recovery from disturbance. For marine reserves to protect biodiversity and enhance populations of species in fished areas, they must be able to sustain focal species (particularly fishery species) within their boundaries, and be spaced such that they can function as mutually replenishing networks whilst providing recruitment subsidies to fished areas. Thus the configuration (size, spacing and location) of individual reserves within a network should be informed by larval dispersal and movement patterns of the species for which protection is required. In the past, empirical data regarding larval dispersal and movement patterns of adults and juveniles of many tropical marine species have been unavailable or inaccessible to practitioners responsible for marine reserve design. Recent empirical studies using new technologies have also provided fresh insights into movement patterns of many species and redefined our understanding of connectivity among populations through larval dispersal. Our review of movement patterns of 34 families (210 species) of coral reef fishes demonstrates that movement patterns (home ranges, ontogenetic shifts and spawning migrations) vary among and within species, and are influenced by a range of factors (e.g. size, sex, behaviour, density, habitat characteristics, season, tide and time of day). Some species move <0.1–0.5 km (e.g. damselfishes, butterflyfishes and angelfishes), <0.5–3 km (e.g. most parrotfishes, goatfishes and surgeonfishes) or 3–10 km (e.g. large parrotfishes and wrasses), while others move tens to hundreds (e.g. some groupers, emperors, snappers and jacks) or thousands of kilometres (e.g. some sharks and tuna). Larval dispersal distances tend to be <5–15 km, and self‐recruitment is common. Synthesising this information allows us, for the first time, to provide species, specific advice on the size, spacing and location of marine reserves in tropical marine ecosystems to maximise benefits for conservation and fisheries management for a range of taxa. We recommend that: (i) marine reserves should be more than twice the size of the home range of focal species (in all directions), thus marine reserves of various sizes will be required depending on which species require protection, how far they move, and if other effective protection is in place outside reserves; (ii) reserve spacing should be <15 km, with smaller reserves spaced more closely; and (iii) marine reserves should include habitats that are critical to the life history of focal species (e.g. home ranges, nursery grounds, migration corridors and spawning aggregations), and be located to accommodate movement patterns among these. We also provide practical advice for practitioners on how to use this information to design, evaluate and monitor the effectiveness of marine reserve networks within broader ecological, socioeconomic and management contexts.     

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.     

Marine reserves indirectly affect fine‐scale habitat associations,but not overall densities,of small benthic fishes

Many large, fishery‐targeted predatory species have attained very high relative densities as a direct result of protection by no‐take marine reserves. Indirect effects, via interactions with targeted species, may also occur for species that are not themselves targeted by fishing. In some temperate rocky reef ecosystems, indirect effects have caused profound changes in community structure, notably the restoration of predator–urchin–macroalgae trophic cascades. Yet, indirect effects on small benthic reef fishes remain poorly understood, perhaps because of behavioral associations with complex, refuge‐providing habitats. Few, if any, studies have evaluated any potential effects of marine reserves on habitat associations in small benthic fishes. We surveyed densities of small benthic fishes, including some endemic species of triplefin (Tripterygiidae), along with fine‐scale habitat features in kelp forests on rocky reefs in and around multiple marine reserves in northern New Zealand over 3 years. Bayesian generalized linear mixed models were used to evaluate evidence for (1) main effects of marine reserve protection, (2) associations with habitat gradients, including complexity, and (3) differences in habitat associations inside versus outside reserves. No evidence of overall main effects of marine reserves on species richness or densities of fishes was found. Both richness and densities showed strong associations with gradients in habitat features, particularly habitat complexity. In addition, some species exhibited reserve‐by‐habitat interactions, having different associations with habitat gradients inside versus outside marine reserves. Two species (Ruanoho whero and Forsterygion flavonigrum) showed stronger positive associations with habitat complexity inside reserves. These results are consistent with the presence of a behavioral risk effect, whereby prey fishes are more strongly attracted to habitats that provide refuge from predation in areas where predators are more abundant. This work highlights the importance of habitat structure and the potential for fishing to affect behavioral interactions and the interspecific dynamic attributes of community structure beyond simple predator–prey consumption and archetypal trophic cascades.     

Evidence of artisanal fishing impacts and depth refuge in assemblages of Fijian reef fish

Protection from fishing generally results in an increase in the abundance and biomass of species targeted by fisheries within marine reserve boundaries. Natural refuges such as depth may also protect such species, yet few studies in the Indo Pacific have investigated the effects of depth concomitant with marine reserves. We studied the effects of artisanal fishing and depth on reef fish assemblages in the Kubulau District of Vanua Levu Island, Fiji, using baited remote underwater stereo-video systems. Video samples were collected from shallow (5–8 m) and deep (25–30 m) sites inside and outside of a large old marine reserve (60.6 km², 13 years old) and a small new marine reserve (4.25 km², 4 years old). Species richness tended to be greater in the shallow waters of the large old reserve when compared to fished areas. In the deeper waters, species richness appeared to be comparable. The difference in shallow waters was driven by species targeted by fisheries, indicative of a depth refuge effect. In contrast, differences in the abundance composition of the fish assemblage existed between protected and fished areas for deep sites, but not shallow. Fish species targeted by local fisheries were 89% more abundant inside the large old reserve than surrounding fished areas, while non-targeted species were comparable. We observed no difference in the species richness or abundance of species targeted by fisheries inside and outside of the small new reserve. This study suggests that artisanal fishing impacts on the abundance and species richness of coral reef fish assemblages and effects of protection are more apparent with large reserves that have been established for a long period of time. Observed effects of protection also vary with depth, highlighting the importance of explicitly incorporating multiple depth strata in studies of marine reserves.     

Marine reserve design theory for species with ontogenetic migration

Models for marine reserve design have been developed primarily with ‘reef fish’ life histories in mind: sedentary adults in patches connected by larval dispersal. However, many fished species undertake ontogenetic migrations, such as from nursery grounds to adult spawning habitats, and current theory does not fully address the range of reserve options posed by that situation. I modelled a generic species with ontogenetic migration to investigate the possible benefits of reserves under three alternative scenarios. First, the fishery targets adult habitat, and reserves can sustain yields under high exploitation, unless habitat patches are well connected. Second, the fishery targets the nursery, and reserves are highly effective, regardless of connectivity patterns. Third, the fishery targets both habitats, and reserves only succeed if paired on adjacent, well-connected nursery and adult patches. In all cases, reserves can buffer populations against overexploitation but would not enhance fishery yield beyond that achievable by management without reserves. These results summarize the general situations in which management using reserves could be useful for ontogenetically migrating species, and the type of connectivity data needed to inform reserve design.     

On the Optimal Size of Marine Reserves

The excessive and unsustainable exploitation of our marine resources has led to the promotion of marine reserves as a fisheries management tool. Marine reserves, areas in which fishing is restricted or prohibited, can offer opportunities for the recovery of exploited stock and fishery enhancement. This study examines the impact of the creation of marine protected areas, from both economic and biological perspectives. The consequences of reserve establishment on the long-run equilibrium fish biomass and fishery catch levels are evaluated. We include reserve size as control variable to maximize catch at equilibrium. A continuous time model is used to simulate the effects of reserve size on fishing catch. Fish movements between the sites is assumed to take place at a faster time scale than the variation of the stock and the change of the fleet size. We take advantage of these two time scales to derive a reduced model governing the dynamics of the total fish stock and the fishing effort. Simulation results suggest that the establishment of a protected marine reserve will always lead to an increase in total fish biomass, an optimal size of a marine reserve can achieve to maximize the catch at equilibrium.     

Life-history changes in exploited reef fishes on the east coast of South Africa

Synopsis The impact of exploitation on various life-history characteristics of two sex changing, reef-dwelling sparid species was examined by comparing populations protected in a large marine reserve with those adjacent to the reserve. Like other sparids, Chrysoblephus laticeps and C. cristiceps grow slowly and are long lived, reaching ages of 17 and 21 years, respectively. No significant differences in the growth rate of C. laticeps were measured, but growth in C. cristiceps was significantly slower in the exploited population. Observed data showed that sex ratios outside the marine reserve were skewed towards the females, a result of size selective exploitation. Size at sex change was also significantly smaller for C. cristiceps in the exploited area, but not so for C. laticeps. This difference between the species was explained as a function of the size at recruitment into the fishery and the degree of protection afforded both large females and male fish. Considering the possibility that reproduction could be impaired as a result of changes in population structure, the tactic of protection through marine reserves is supported as a hedge against recruitment failure.     

Changes in Fish Assemblages following the Establishment of a Network of No-Take Marine Reserves and Partially-Protected Areas

Networks of no-take marine reserves and partially-protected areas (with limited fishing) are being increasingly promoted as a means of conserving biodiversity. We examined changes in fish assemblages across a network of marine reserves and two different types of partially-protected areas within a marine park over the first 5 years of its establishment. We used Baited Remote Underwater Video (BRUV) to quantify fish communities on rocky reefs at 20–40 m depth between 2008–2011. Each year, we sampled 12 sites in 6 no-take marine reserves and 12 sites in two types of partially-protected areas with contrasting levels of protection (n = 4 BRUV stations per site). Fish abundances were 38% greater across the network of marine reserves compared to the partially-protected areas, although not all individual reserves performed equally. Compliance actions were positively associated with marine reserve responses, while reserve size had no apparent relationship with reserve performance after 5 years. The richness and abundance of fishes did not consistently differ between the two types of partially-protected areas. There was, therefore, no evidence that the more regulated partially-protected areas had additional conservation benefits for reef fish assemblages. Overall, our results demonstrate conservation benefits to fish assemblages from a newly established network of temperate marine reserves. They also show that ecological monitoring can contribute to adaptive management of newly established marine reserve networks, but the extent of this contribution is limited by the rate of change in marine communities in response to protection.     

Changes in fish assemblages following 10 years of protection in Tasmanian marine protected areas

Most studies examining effects of marine protected areas (MPAs) on fish assemblages are potentially confounded, either because they are once off comparisons between fished and unfished locations, or because they are snapshot studies over a fixed period. Here we compare long-term changes within fully protected Tasmanian marine reserves with changes at external reference sites on an annual basis over the first ten years of protection. The results highlight the importance of long-term datasets for differentiating changes occurring over differing time scales. Notable results include a statistically significant increase in abundance of Latridopsis forsteri and large fish (> 300 mm) when examined across all reserves relative to controls, and a 10-fold increase in the abundance of large fish and a doubling of per site species richness of large fish within the Tinderbox Marine Reserve relative to controls. Short-term resident species that recruit sporadically show very different patterns in reserves compared to those that recruit regularly and have long-term age-class storage. While several recent reviews have suggested size of MPAs and duration of protection has little influence on the extent of recovery, our results suggest this is not the case and that responses can be slow, complex and species-specific. The extent of localised fishing pressure appeared to have a substantial influence on the degree of change detected, potentially confounding meta-analyses of recovery rates in MPAs if overlooked as a relevant parameter.     

Opening of an MPA to fishing: Natural variations in the structure of a coral reef fish assemblage obscure changes due to fishing

Marine Protected Areas (MPAs) are known to enhance diversity, density and biomass of coral reef fishes and to modify the size and trophic structures of these fish assemblages. Opening to fishing has the opposite effects, but on a much shorter time scale. The present study compares the evolution of the fish assemblages of two adjacent reef zones, both initially MPAs, one of them being afterwards opened to fishing. The study was conducted on Aboré Reef, a New Caledonian barrier reef (SW Pacific) which constituted a 148 km² marine protected area, of which 69 km² are within the lagoon. Two surveys of a coral reef fish assemblage, using underwater visual censuses, were performed, the first one was conducted in July 1993 following 5 years of protection from fishing, the second one was conducted in July 1995; part of the reef having been opened to fishing activity in September 1993. This study examined the effects of two factors on these fish communities: time (1993 vs. 1995) and zone (reefs protected from fishing vs. unprotected reefs); the interaction of these two factors indicating an effect of either protection or opening to fishing. Diversity (species/transect), density and biomass were tested for all species together (377 species), then according to diet, size and commercial use. There was a significant decrease over time of most values in both fished and unfished areas, the decrease being greater in the zone opened to fishing. The magnitude of decrease over time was within the range of known time variations from other studies in New Caledonia and other Pacific locations. However, this decrease was so large that it prevented the detection of effects linked to protection. Only some minor effects could be detected for 16 species with no specific pattern according to diet, size or use. The density and biomass of species of low commercial value were also affected by opening to fishing. Relative changes in diversity could be better detected than relative changes in density or biomass. This study demonstrates that on a short-term basis (2 years), natural variations can be of larger magnitude than changes that may be induced by management options, especially when fishing pressure is not very high.     

基于熵权的珠江三角洲自然保护区综合评价

运用熵权法对珠江三角洲自然保护区进行了综合评价。结果表明:(1)48个样本的保护区中7个为高水平,30个为较高水平,10个为一般,1个较差。从总体水平上看,虽然具有一部分较高水平的保护区,但珠江三角洲的保护区建设水平存在明显的差异,深圳、中山、佛山等地亟待建立新的保护区加以补充;(2)保护区的综合水平与面积具有明显的相关关系,建立了综合水平与面积之间的回归模型,并进一步求出珠江三角洲生态类型自然保护区理论上的最适宜面积;(3)区域保护区综合水平的提高与最大面积指数正相关,区域可以通过建设面积相对较大的优质保护区成为区域生态增长极,通过物种的交流和环境影响拉动其它保护区的发展,但区域范围内保护区之间面积差异不能太大;(4)自然干扰对沿海及岛屿类型的保护区综合水平影响相对较大。     

Spatial and temporal variation of abundance,biomass and diversity within marine reserves in the Philippines

Aim The objective of this study was to investigate the influence of protection duration (years of fishing closure) and location (distance from shore) on reef fish diversity. Location Danajon Double Barrier Reef, Bohol, Philippines. Methods Reef fish abundance and size structure, by species, were obtained monthly using replicated underwater visual belt transects (n = 8; 70 × 5‐m belt transects) over 3 years (2002–2005) at eight sites that included six marine reserves and two unprotected reef areas. We analysed species accumulation curves, diversity indices and abundance–biomass comparison (ABC) curves within and across the study sites to assess the influence of protection duration and location. Results Analyses showed that longer protection duration impacted reef fish diversity at both inshore and offshore sites by shifting ABC curves from higher abundance than biomass curves at fished sites to higher biomass than abundance curves at most of the protected sites. Protection duration did not significantly influence either the rate of species accumulation within sites or the 12 diversity indices measured across the study sites. The offshore sites consistently showed higher rates of species accumulation and diversity indices values than inshore sites with similar protection duration. One protected offshore young marine reserve site that has been assessed as the least well‐managed showed patterns more consistent with the fished sites. Main conclusions Analyses showed that protection duration mainly impacted diversity by increasing the dominance of large‐bodied species and enhancing total biomass. Besides protection duration, reserve location influenced species accumulation curves and diversity indices.     

Catastrophes,connectivity and Allee effects in the design of marine reserve networks

Catastrophic events, like oil spills and hurricanes, occur in many marine systems. One potential role of marine reserves is buffering populations against disturbances, including the potential for disturbance-driven population collapses under Allee effects. This buffering capacity depends on reserves in a network providing rescue effects, setting up a tradeoff where reserves need to be connected to facilitate rescue, but also distributed in space to prevent simultaneous extinction. We use a set of population models to examine how dispersal ability and the disturbance regime interact to determine the optimal reserve spacing. We incorporate fishing in a spatially-explicit model to understand the effect of objective choice (e.g. conservation versus fisheries yield) on the optimal reserve spacing. We show that the optimal spacing between reserves increases when accounting for catastrophes with larger spacing needed when Allee effects interact with catastrophes to increase the probability of extinction. We also show that classic tradeoffs between conservation and fishing objectives disappear in the presence of catastrophes. Specifically, we found that at intermediate levels of disturbance, it is optimal to spread out reserves in order to increase both population persistence and to maximize spillover into non-reserve areas.     

Effects of marine reserves versus nursery habitat availability on structure of reef fish communities

No-take marine fishery reserves sustain commercial stocks by acting as buffers against overexploitation and enhancing fishery catches in adjacent areas through spillover. Likewise, nursery habitats such as mangroves enhance populations of some species in adjacent habitats. However, there is lack of understanding of the magnitude of stock enhancement and the effects on community structure when both protection from fishing and access to nurseries concurrently act as drivers of fish population dynamics. In this study we test the separate as well as interactive effects of marine reserves and nursery habitat proximity on structure and abundance of coral reef fish communities. Reserves had no effect on fish community composition, while proximity to nursery habitat only had a significant effect on community structure of species that use mangroves or seagrass beds as nurseries. In terms of reef fish biomass, proximity to nursery habitat by far outweighed (biomass 249% higher than that in areas with no nursery access) the effects of protection from fishing in reserves (biomass 21% lower than non-reserve areas) for small nursery fish (≤ 25 cm total length). For large-bodied individuals of nursery species (>25 cm total length), an additive effect was present for these two factors, although fish benefited more from fishing protection (203% higher biomass) than from proximity to nurseries (139% higher). The magnitude of elevated biomass for small fish on coral reefs due to proximity to nurseries was such that nursery habitats seem able to overrule the usually positive effects on fish biomass by reef reserves. As a result, conservation of nursery habitats gains importance and more consideration should be given to the ecological processes that occur along nursery-reef boundaries that connect neighboring ecosystems.     

Go with the flow: tidal import and export of larvae from semi-enclosed bays

There is a growing body of evidence that suggests the effective functioning of marine reserves is dependent on the dispersal and recruitment of larvae. Enhanced production inside reserves is predicted to lead to a net larval export and increased settlement and recruitment outside reserve boundaries. However, larval retention in bays is also well documented. Since bays are increasingly being used as reserve areas, planktonic larvae of benthic marine invertebrates were sampled from two semi-enclosed marine reserves during flood and ebb tides to determine whether these bays are acting as net exporters of larvae. Neither reserve was a net importer or exporter of species richness, larval abundance or diversity, although one reserve showed a small export of species richness during the hours of darkness. Both reserves balanced the net import of some species with a net export of others, which was generally related to adult or larval abundance, although exceptions were found in one reserve. Significant effects of light were found, with the net import or export of some species occurring exclusively during either the hours of daylight or darkness. An increased understanding of larval sink-source dynamics in bays is essential for ensuring their effective use as marine reserves to meet specific conservation needs. Guest editors: J. Davenport, G. Burnell, T. Cross, M. Emmerson, R. McAllen, R. Ramsay & E. Rogan Challenges to Marine Ecosystems     

Trajectories and magnitude of change in coral reef fish populations in Philippine marine reserves: a meta-analysis

Marine reserves are widely implemented worldwide to meet both conservation and fisheries management goals. This study examines the efficacy of Philippine marine reserves using meta-analysis by comparing variations in fish density (1) between reserves and adjacent fished reefs (spatial comparison), (2) within reserves before establishment relative to years following the establishment (temporal comparison), and (3) among reserves classified based on size, age, and enforcement capacity. A grand (total) mean of nineteen 22.3 ha coral reef reserves, protected for a mean duration of 8.2 years, were included in the meta-analyses. The overall density of fishes was higher in the reserves compared with the fished reefs and this difference was largely accounted for by exploited fishes. However, the overall density of fishes within the same reserves remained similar from the period before its establishment to several years following its establishment. Only the density of nonexploited fishes increased significantly during years subsequent to the establishment of the reserves. Neither age nor size of reserves correlated with pattern of change in fish density following the establishment of the reserves; however, fish density was consistently higher in larger and older reserves relative to smaller and younger reserves in the spatial comparison. Furthermore, well-enforced reserves had higher density of exploited fishes relative to less-enforced reserves in both spatial and temporal comparisons. In general, the magnitude and trajectory of change in fish density following the establishment of Philippine marine reserves are influenced by (1) functional groups of fishes under consideration, (2) size and age of the reserve, and (3) level of enforcement of the regulatory mechanisms necessary to sustain a marine reserve.     

53 A Seaweed's perspective on marine reserves

In response to major changes in coastal ecosystems in recent decades, a number of governmental agencies around the world are establishing marine reserves – areas where removal of animals or plants is prohibited. Although marine reserves are touted as an ecosystem based approach to management of marine resources, the vast majority of attention on reserve design and impact focuses solely on fish. Although a few species of algae are commercially harvested, most are not. As a result, they will receive little direct benefit from protection by reserves aside from habitat protection. From the perspective of a seaweed, the primary impacts of marine reserves will therefore be indirect through species interactions. We examine the rapidly growing theoretical and empirical literature on marine reserves to anticipate the likely responses of seaweeds to exclusion of fishing. The key issues that emerge are: the trophic level of prior fishing and the dispersal scales of seaweeds relative to their competitors and consumers. The latter issue is poorly understood and poses a key challenge to phycologists if we are to effectively incorporate seaweeds into future marine reserve design.     

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.