Marine reserve benefits and recreational fishing yields: The winners and the losers

Marine reserves constitute effective tools for preserving fish stocks and associated human benefits. However, not all reserves perform equally, and predicting the response of marine communities to management actions in the long run is challenging. Our decadal-scale survey of recreational fishing yields at France’s 45-year old Cerbère-Banyuls marine reserve indicated significant protection benefits, with 40–50% higher fishing yields per unit effort in the partial-protection zone of the reserve (where fishing is permitted but at a lower level) than in surrounding non-reserve areas. Over the period 2005–2014, catch per unit effort (CPUE) declined both inside and outside the reserve, while weight per unit effort (WPUE) increased by 131% inside and decreased by 60% outside. Different CPUE and WPUE trajectories among fish families indicated changing catch assemblages, with yields increasing for the family most valued by fisheries, Sparidae (the ecological winners). However, reserve benefits were restricted to off-shore fishermen (the social winners), as on-shore yields were ~4 times lower and declining, even inside the reserve. Our study illustrates how surveys of recreational fishing yields can help evaluate the effectiveness of marine protected areas for key social and ecological protagonists. We show that, more than four decades after its establishment, fishing efficiencies at the historical Cerbère-Banyuls marine reserve are still changing, but benefits in terms of catch abundance, weight, and composition remain predominantly restricted to off-shore fishermen. Further regulations appear necessary to guarantee that conservation strategies equitably benefit societal groups.

buffer zone of partial protection surrounding the fully protected no-take area (Fig 1). In contrast, 90 no restrictions apply outside of the reserve where fishing follows the French national regulation 91 for the Mediterranean Sea. Therefore, we hypothesized that fishing yields would differ between 92 the partial protection zone of the reserve, which benefits from the vicinity of the fully protected   We used three biological indicators commonly used to characterize fishing yields: catch 111 abundance, weight, and composition. We tested for differences in catch per unit effort (CPUE) 112 and weight per unit effort (WPUE) between fishermen fishing inside versus outside of the 113 reserve, as well as on-shore along the coastline versus off-shore from boats. CPUE and WPUE  Table). Based on an unprecedented survey of recreational fishing activities in the Mediterranean, 119 our study provides a decadal-scale evaluation of the effectiveness of the Cerbère-Banyuls nature   Table). The reserve also hosts  the reserve and 988 in surrounding areas (Fig 1). All interviews took place during daytime, and 212 targeted randomly-selected recreational anglers on-shore along the beaches, jetties, and rocky 213 coastline (650 interviews), and off-shore onboard small, typically 4-7 m boats (831 interviews).

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Interviews were conducted anonymously after informed consent for study participation from 215 each subject.

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Statistical analyses 218 We used generalized linear models to evaluate spatio-temporal variability in CPUE and 219 WPUE between inside and outside the reserve, for fishermen fishing on-and off-shore, for all  Table). For clarity 224 and ease of narration, changes in fishing yields are described sequentially, from the main effects 225 of the reserve and time alone (Fig 2), to the additional effects of fishermen groups (Fig 3) and 226 fish families (Fig 4). Preliminary tests of deviance of model residuals indicated a negative-

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All species catches 234 Over the course of the study, a total of 5,864 individual catches for a fishing effort of  Table).  The effects of the reserve on CPUE and WPUE trajectories also differed among the two 255 fishermen groups (Fig 3, S3 Table). On-shore, fishing yield was in decline both in the reserve (-   (Fig 4a) was in decline on-shore both inside  Table). WPUE Spar showed a different pattern over this period (Fig 4b); 289 on-shore values increased in the reserve (+99%, from 34.2 to 68.0 g.line -1 .h -1 ) but decreased in 290 surrounding non-reserve areas (-93%, from 177.4 to 12.8 g.line -1 .h -1 ), and off-shore values 291 increased both in the reserve (+78%, from 107.4 to 191.3 g.line -1 .h -1 ) and in nearby non-reserve 292 waters (+224%, from 70.8 to 229.5 g.line -1 .h -1 ). Overall, contrasting WPUE Spar trajectories were 293 observed between fishermen performing on-versus off-shore, independently from being located 294 inside or outside of the reserve (p=0.0013, S2 Table).    Table).

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While off-shore fishing is restricted to boat users and segregates fishing effort in a two-372 dimensional space throughout the reserve, the near-shore is potentially accessible to all 373 fishermen and concentrates fishing pressure on a few accessible sites (mostly along beeches and 374 jetties) along the mono-dimensional stretch of the coastline (Fig 1). Moreover, near-shore 375 habitats are typically more exposed to other forms of degradation that impact marine biota, these new measures on fishing efforts and yields remain to be evaluated, the number of fishing 387 permits could further be reduced in the future as annual user-permit demands have been below 388 the 1,000 threshold in recent years. Nevertheless, there is growing concern that declining fishing 389 yields could jeopardize the popularity of recreational fishing, an emblematic activity in the 390 region, with significant economic consequences for the associated sectors including bait markets, 391 harbors, and tourism. Sparidae, but were 2-3 times higher in the reserve for Serranidae. Similarly, equivalent levels of 398 catch abundance and weight were found on-and off-shore for Labridae, while yields were twice 399 higher off-shore for Sparidae and ~20 times higher for Serranidae. The heterogeneity of benthic 400 habitats was previously found to influence spatial variability in fish assemblage abundance and 401 composition more strongly than protection status at the Cerbère-Banyuls marine reserve (Claudet  Table). As such, the increasing yields recorded for Sparidae indicate the reserve was effective in 414 supporting catches of large individuals from species of high-value to fisheries within the 415 protected area as well as an apparent spillover benefit to adjacent off-shore areas as expected for  Table)  progressive transfer in catch biomass in space from on-shore to off-shore, as well as in 426 composition from smaller, less-targeted fish to larger species that are of higher value to fisheries.

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Implications for management 429 Our study shows that surveys of recreational fishing activities can constitute robust  Table), recreational fishing yields at the Cerbère-Banyuls marine reserve are  We would like to thank all the recreational fishermen who accepted to participate in our 471 interviews. We also thank F. Huguet, C. Pou, C. Marjorie, F. Mirbeau, J. Jean-Baptiste, R. providing us port space. We are also grateful to J. Ballard for improvements on the manuscript.