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.

Introduction

Despite increasing management efforts, the decline of fishing yields remains a global concern [1]. This is especially true in the Mediterranean Sea where human impacts to the marine environment are diverse, intense, and increasing [2]. Indeed, the Mediterranean is considered the most overfished marine basin on the planet and poses severe management challenges, as exploitation of marine resources in this nearly-enclosed sea is shared among 21 bordering countries whose economic development is tied to activities impinging on the marine environment [3-5]. While Mediterranean marine biota are threatened by a multitude of anthropogenic stressors including pollution and eutrophication, climate change, invasive species, marine transport, aquaculture, and tourism, the major historical and current drivers of declining biodiversity and productivity are habitat loss and fishing [2, 6].

Fish landings in the Mediterranean have been decreasing since the mid-1980’s, despite expanding fishing efforts toward lower trophic levels and the deeper sea [1, 7, 8]. This decreasing catch rate has resulted in declines of commercial fishing activities. In contrast, recreational fishing has been on the rise, particularly along the European coast of the Mediterranean [3, 4, 9]. As in many other regions, the relative contributions of commercial versus recreational fishing to the local socio-economy and decline of fish stocks have yet to be quantified throughout the Mediterranean [8, 10–12], though recreational fishing is suspected to exert a strong and increasing pressure, particularly on highly targeted marine species [9, 13–16]. In the Mediterranean and elsewhere, strategies for preserving fish stocks consist primarily of regulating fishing efforts through gear restrictions (gear type and number), fishing yields through quotas (catch size and bag limits), and fishing areas and seasons through exclusion zones and marine reserves [6, 10]. However, the long term effectiveness of these management strategies for preserving species abundance and ecosystem services is difficult to predict [17, 18]. In the face of such uncertainties, the preservation of fisheries resources and associated socio-economic benefits poses serious regulatory challenges in terms of implementing appropriate measures for resource durability and equitable access [3, 19]. In this context, identifying social and ecological protagonists vulnerable to environmental decline can refine regulatory strategies and help define win-win, sustainable management for people and ecosystems [20-22].

We performed a decadal scale survey (2005–2014) of recreational fishing activities at the Cerbère-Banyuls marine reserve (Fig 1), one of the oldest marine protected areas in the Mediterranean, to evaluate the effectiveness of local management efforts in preserving fisheries resources. The survey consisted of ~1,500 on-site interviews with recreational fishermen fishing inside and outside the reserve, and recorded ~6,000 individual catches representing a total weight of ~1 ton for a fishing effort of ~5000 line-hours. Within the reserve, fishing is subject to restrictions (gear restrictions and bag and size limits, see section 2.1.) and only takes place in a buffer zone of partial protection surrounding the fully protected no-take area (Fig 1). In contrast, no restrictions apply outside of the reserve where fishing follows the French national regulation for the Mediterranean Sea. Therefore, we hypothesized that fishing yields would differ between the partial protection zone of the reserve, which benefits from the vicinity of the fully protected no-take area and undergoes restricted fishing pressure, as compared to surrounding no-reserve areas. Similarly, because reserve benefits are often not equally distributed in space and among species undergoing different fishing pressure [17, 23–26], we also hypothesized that reserve benefits would differ for fishermen fishing on-shore (access limited to the coastline of the reserve) and off-shore (unrestricted boat access to the entire reserve), as well as among fish families differently targeted by fisheries.

Fig 1

Map of the study area indicating the position of the Cerbère-Banyuls marine reserve’s fully protected core area (no-take zone), buffer zone of partial protection (fishing allowed with restrictions, see Methods), and control outside area (no specific regulation on fishing) in relation to the towns of Banyuls-sur-Mer and Cerbère and the two capes, Cap Béar and Cap Cerbère.

Established in 1974, Cerbère-Banyuls is one of the oldest marine protected areas of the Mediterranean Sea. The arrow in the insert indicates the position of the reserve in the natural marine park of the Gulf of Lion situated in the north-western corner of the Mediterranean, at the border between France and Spain. Isobaths indicate depth variation every 10 meters. Maps were produced using the open source program QGIS.

We used three biological indicators commonly used to characterize fishing yields: catch abundance, weight, and composition. We tested for differences in catch per unit effort (CPUE) and weight per unit effort (WPUE) between fishermen fishing inside versus outside of the reserve, as well as on-shore along the coastline versus off-shore from boats. CPUE and WPUE trajectories were also compared among the three dominant fish families (S1 Table), namely Sparidae (sea breams), Serranidae (groupers), and Labridae (wrasses), each exhibiting different levels of species diversity, occupying different positions in habitats and trophic levels, and with different values for fisheries [3, 9, 27]. Based on an unprecedented survey of recreational fishing activities in the Mediterranean, our study provides a decadal-scale evaluation of the effectiveness of the Cerbère-Banyuls nature reserve, one of the most preserved marine reserves in the region (see Methods), for supporting fishing yields. Our results shed light on the consequences of fishing regulations for key social and ecological protagonists with implications for adaptive management of fishery resources.

Methods

Ethics statement

This study involved interviews with recreational fishermen. The interviews were conducted anonymously after informed consent for study participation from each subject. The survey methodology and material was approved by the University of Perpignan.

The Cerbère-Banyuls natural marine reserve

The Cerbère-Banyuls natural marine reserve (www.reserves-naturelles.org/cerbere-banyuls) is a French marine protected area situated in the north-western corner of the Mediterranean Sea (Fig 1). Established in 1974, it is one of the oldest marine reserves in the Mediterranean and is managed by the Departmental Council of the Pyrénées-Orientales. With over 100,000 annual visitors, including more than 30,000 scuba-divers in recent years, the Cerbère-Banyuls marine reserve contributes greatly to the region’s community character and socio-economic development [11, 24]. Thanks to its ecological wealth and management, the reserve is recognized since 2014 as one of the 40 sites listed on IUCN’s Green List of Protected Natural Areas (www.iucn.org), and is since 2018 among the 16 Blue Parks distinguished as outstanding marine protected areas by the Marine Conservation Institute (https://globaloceanrefuge.org). Since 2011, the reserve is part of the larger, 4,010 km2 in area natural marine park of the Gulf of Lion (www.parc-marin-golfe-lion.fr). This is the largest marine park in the Mediterranean Sea, and is managed by the French Agency for Biodiversity. At this stage, there are no restrictions regarding fishing and other human usages in the park, though scientists, managers, and representatives regularly hold discussions through committees, workshops and ongoing projects to deliberate on future plans.

The reserve comprises a small nucleus of 65 ha of fully protected area, where only recreational navigation, surface swimming, and scientific diving are authorized (Fig 1). This area is surrounded by a larger, 650 ha buffer zone of partial protection where recreational activities such as scuba-diving, boating, and daytime angling are authorized, but subject to quotas and restrictions that are stricter than the general French coastal fishing regulations for the Mediterranean Sea which apply outside of the reserve [28-30]. During the studied period, anglers in the reserve were restricted to a maximum of 2 lines with up to 6 hooks if fishing on-shore, and 12 hooks if fishing from a boat. No restrictions on lines and hooks applied outside of the reserve. The number of recreational fishermen fishing in the reserve is regulated by a free but mandatory annual permit; up to 1,500 permits were issued annually over the course of this study. Species-specific minimum catch sizes and maximum bag limits also apply, and spearfishing is forbidden within the reserve [29]. Some commercial fishing also takes place in the partial protection zone of the reserve, with a fleet of 4–15 artisanal boats registered annually during the studied period [11]. Surveys of recreational and commercial fishing indicate that catches around the Cerbère-Banyuls marine reserve consist primarily of fish belonging to the families Sparidae, Serranidae, and Labridae [28, 30] (S1 Table). The reserve also hosts recovering populations of dusky grouper (Epinephelus marginatus, family Serranidae) and brown meagre (Sciaena umbra, family Sciaenidae) which are protected from fishing by national moratoria.

Survey methodology and design

Recreational fishing activities in and around the Cerbère-Banyuls marine reserve were surveyed during four monitoring campaigns performed in 2005, 2009, 2010–2011, and 2013–2014 in an area expanding from Cap Béar in the north to Cap Cerbère in the south and with a water depth ranging 0–90 m (Fig 1). Recreational fishing refers here to all non-commercial fishing activities that are carried out mainly for leisure, where catches are either used for personal consumption, offered to family or friends, or released; the sale of recreational fishing yield being illegal by definition [3]. It encompasses multiple forms of activities performed on- and off-shore, mainly angling, trolling, spearfishing, and shellfish gathering [9, 15, 16].

Surveys consisted of on-site interviews with fishermen in a roving creel survey design [31]. Fishing gear and effort (number and type of lines and hooks, fishing duration, etc.) as well as catch abundance, composition, and size were recorded, including of discarded specimen. The survey instrument was a structured interview featuring a standardized list of questions asked to each participant [30]. Among the multiple approaches that can be used to quantify fishing yields (e.g. scientific campaigns, fisheries logbooks, telephonic surveys) [14, 15, 32], on-site interviews with recreational fishermen have the advantage of maximizing data acquisition in time and space while supporting robust data quality via direct observation of social and ecological descriptors of fisheries (e.g. fishermen abundance, fishing efforts and techniques, catch characteristics). This form of participatory science also promotes positive interactions with fishermen via frequent contact with users (e.g. for increasing awareness and building trust in management strategies). Limitations of such interview-based approaches include the dependency of data quality on user responses to questionnaires. Although the proportion of unreported catch is difficult to evaluate, our interviews indicated that many local fishermen recognized the role of the reserve in preserving marine resources, and it is likely that the majority were honest in their responses. Nevertheless, we assumed the proportion of unreported catch to be relatively constant over time, with no implication on the dynamics of fishing yields as quantified in our study.

Our surveys specifically targeted anglers, who constitute the largest proportion of the local recreational fishing population, fish throughout the year, and are easy to approach for interviews. Fishing gear commonly used by anglers in the study area consist primarily of lures and baited hooks mounted on lines thrown by hand or rod and equipped with weights or floaters [30]. Catch sizes were measured when possible, or otherwise estimated visually or based on fishermen’s declarations (e.g. for discarded yields). Catch weights were estimated using length-weight relationships of species from the literature (www.fishbase.org) with locally estimated parameters when available [33]. Fishing efforts (in line.hour) were calculated based on the number of fishing lines and hooks used by each fishermen, and the time spent between when the fishermen declared starting fishing and the interview. Fishing yield was quantified by calculating catch per unit effort (CPUE, number per line per hour) and weight per unit effort (WPUE, gram per line per hour) for all species combined, as well as individually for the three major fish families recorded (Sparidae, Serranidae, Labridae) [34] which represented >85% of catches in number and >65% of the overall weight captured (see S1 Table for a list of the species recorded for each family). CPUE and WPUE are standard metrics of fishing yields per unit of effort, facilitating comparisons of efficiency among different fishing techniques, targets, and regulations [25, 32, 35].

A total of 1,481 interviews were performed between 2005 and 2014, including 493 within the reserve and 988 in surrounding areas (Fig 1). All interviews took place during daytime, and targeted randomly-selected recreational anglers on-shore along the beaches, jetties, and rocky coastline (650 interviews), and off-shore onboard small, typically 4–7 m boats (831 interviews). Interviews were conducted anonymously after informed consent for study participation from each subject.

Statistical analyses

We used generalized linear models [36] to characterize differences in CPUE and WPUE trajectories between inside and outside the reserve, for fishermen fishing on- and off-shore, for all species catches and each of the 3 major fish families (Sparidae, Serranidae, Labridae). Three-factor models were initially designed to test for differences in yield trajectories (separately for CPUE and WPUE response variables) in time (continuous explanatory variable Time: 2005–2014), space (categorical explanatory variable Reserve: in vs out), among fishermen groups (categorical explanatory variable Fishermen: on- vs off-shore), and their interactions (CPUE ~ Time × Reserve × Fishermen, WPUE ~ Time × Reserve × Fishermen). However, because some of the models did not converge due to over-parametrization [37], simpler two-factor models (Time × Reserve) were used to characterize CPUE and WPUE trajectories separately for on- and off-shore fishermen. Similar results were found when CPUE and WPUE trajectories were characterized using three-factor models (using data from both fishermen groups) and two-factor models (separately per fishermen group). For consistency, only the latter are reported herein (S2 Table). For clarity and ease of narration, changes in fishing yields as expressed in CPUE and WPUE are described sequentially, from the main effects of the reserve and time alone (Fig 2), to the additional effects of fishermen groups (Fig 3) and fish families (Sparidae, Serranidae, Labridae, Fig 4). A separate set of models was used to compare average CPUE and WPUE values over the entire study period (2005–2014), for all catches and by fish family, between fishing zones and fishermen groups (S3 Table). Preliminary tests of deviance of model residuals indicated a negative-binomial distribution of the data. CPUE and WPUE trajectories were therefore estimated using the glm.nb function with a log link for negative-binomial data from the MASS package [38]. All modeling and graphing were coded in R statistical software (R Core Team). We found similar results when considering fishing effort as number of hooks per hour or number of lines per hour (S1 Fig), only the latter being reported below.

Fig 2

Trends in catch per unit effort (CPUE, a) and weight per unit effort (WPUE, b) of recreational fishermen fishing inside (in) and outside (out) the Cerbère-Banyuls marine reserve.

Curves represent mean trajectories estimated by generalized linear models and shadings indicate 95% confidence intervals. The percent changes in mean CPUE and WPUE between the beginning and the end of the study period are provided as text on the plots. See S4 Table for mean and confidence interval values. Refer to S5 Table for parameter estimates.

Fig 3

Trends in catch per unit effort (CPUE, a) and weight per unit effort (WPUE, b) of recreational fishermen fishing from land along the coastline (land) vs at sea from boats (sea), and inside (in) vs outside (out) the Cerbère-Banyuls marine reserve.

Curves represent mean trajectories estimated by generalized linear models and shadings indicate 95% confidence intervals. The percent changes in mean CPUE and WPUE between the beginning and the end of the study period are provided as text on the plots. See S4 Table for mean and confidence interval values. Refer to S5 Table for parameter estimates.

Fig 4

Trends in catch per unit effort (CPUE) and weight per unit effort (WPUE) of recreational fishermen fishing from land along the coastline (land) vs at sea from boats (sea), and inside (in) vs outside (out) the Cerbère-Banyuls marine reserve for each of the 3 major fish families (Sparidae, Serranidae and Labridae).

Note differences in scale in y-axes. Curves represent mean trajectories estimated by generalized linear models and shadings indicate 95% confidence intervals. The percent changes in mean CPUE and WPUE between the beginning and the end of the study period are provided as text on the plots. See S4 Table for mean and confidence interval values. Refer to S5 Table for parameter estimates.

Results

All species catches

Over the course of the study, a total of 5,864 individual catches for a fishing effort of 5,028.9 line-hours, and a total fishing yield of 947.0 kg for a fishing effort of 4,970.7 line-hours, were recorded. The average catch per unit effort (CPUE) for the period 2005–2014 was 1.7 ±1.0 SE ind.line-1.h-1, and the average weight per unit effort (WPUE) was 228.7 ±1.1 SE g.line-1.h-1. Averaging over all years, fishing yield inside the buffer zone of partial protection in the reserve was 1.4 times higher in terms of catch abundance (CPUE = 2.1 ±1.1 SE vs 1.5 ±1.1 SE ind.line-1.h-1, p = 0.002) and 1.5 times higher in weight, as compared with surrounding areas (WPUE = 288.5 ±1.1 SE vs 198.8 ±1.1 SE g.line-1.h-1, p = 0.018; S3 Table). However, throughout the 2005–2014 survey period, CPUE (Fig 2a) declined both inside (-58%, from 2.5 to 1.1 ind.line-1.h-1) and outside the reserve (-66%, from 2.2 to 0.8 ind.line-1.h-1) following a similar pattern to each other, while contrasting WPUE trajectories were observed(Fig 2b), with values increasing in the reserve (+131%, from 222.3 to 514.0 g.line-1.h-1) and decreasing outside (-60%, from 275.3 to 110.1 g.line-1.h-1; S4 Table).

The effects of the reserve on CPUE and WPUE trajectories also differed among the two fishermen groups (Fig 3, S4 Table). On-shore, fishing yield was in decline both in the reserve (-39% in CPUE from 0.6 to 0.4 ind.line-1.h-1, -29% in WPUE from 74.3 to 52.8 g.line-1.h-1) and in surrounding areas (-53% in CPUE from 1.0 to 0.5 ind.line-1.h-1, and -85% in WPUE 202.2 to 29.5 g.line-1.h-1). Off-shore, catch abundance also declined substantially inside (-74% in CPUE from 4.0 to 1.1 ind.line-1.h-1) and outside the reserve (-55% in CPUE from 2.9 to 1.3 ind.line-1.h-1), but average yield in weight showed a milder decline outside (-17% in WPUE from 320.7 to 264.7 g.line-1.h-1) and was increasing in the reserve (+61% in WPUE from 336.7 to 543.4 g.line-1.h-1). Discriminating catches by fishermen groups alone for the ten-year period revealed on average a 3.6 times lower catch abundance (CPUE = 0.7 ±1.1 SE vs 2.5 ±1.1 SE ind.line-1.h-1, p<0.001) and a 3.9 times lower yield in weight (WPUE = 87.9 ±1.1 SE vs 339.5 ±1.1 SE g.line-1.h-1, p<0.001; S3 Table) from land relative to off-shore.

Sparidae

Sparidae represented 32% (1,895 ind.) of the total recorded catch, and 47% (440.6 kg) of the overall yield in weight. An average CPUESpar of 0.4 ±1.1 SE ind.line-1.h-1 and WPUESpar of 93.2 ±1.1 SE g.line-1.h-1 were recorded for 2005–2014 (considering all years). Sparidae catch abundance and weight did not differ significantly between inside and outside the reserve in this period (CPUESpar = 0.4 ±1.1 SE vs 0.5 ±1.1 SE ind.line-1.h-1, p = 0.422; WPUESpar = 100.7 ±1.2 SE vs 89.4 ±1.2 SE g.line-1.h-1, p = 0.641), but were 2 times higher for fishermen off-shore relative to on-shore (CPUESpar = 0.6 ±1.1 SE vs 0.3 ±1.1 SE ind.line-1.h-1, p<0.001; WPUESpar = 119.1 ±1.2 SE vs 60.2 ±1.2 SE g.line-1.h-1, p = 0.006; S3 Table).

Between 2005 and 2014, average CPUESpar (Fig 4a) was in decline on-shore both inside (-73%, from 0.3 to 0.1 ind.line-1.h-1) and outside the reserve (-34%, from 0.4 to 0.2 ind.line-1.h-1), whereas for off-shore fishermen, a comparatively milder decline was observed in the reserve (-14%, from 0.5 to 0.4 ind.line-1.h-1) and values were increasing in surrounding areas (+35%, from 0.6 to 0.7 ind.line-1.h-1; S4 Table). WPUESpar showed a different pattern over this period (Fig 4b); on-shore values increased in the reserve (+99%, from 34.2 to 68.0 g.line-1.h-1) but decreased in surrounding non-reserve areas (-93%, from 177.4 to 12.8 g.line-1.h-1), and off-shore values increased both in the reserve (+78%, from 107.4 to 191.3 g.line-1.h-1) and in nearby non-reserve waters (+224%, from 70.8 to 229.5 g.line-1.h-1). Overall, contrasting WPUESpar trajectories were observed between fishermen performing on- versus off-shore, independently from being located inside or outside of the reserve (p = 0.0013, S2 Table).

Serranidae

Serranidae represented 42% (2,471 ind.) of the total recorded catch, and 17% (157.4 kg) of the overall yield in weight. An average CPUESerr of 0.8 ±1.1 SE ind.line-1.h-1 and WPUESerr of 55.8 ±1.1 SE g.line-1.h-1 were recorded for 2005–2014 (considering all years). Serranidae catch abundance and weight were respectively 2.4 and 3 times higher in the reserve relative to surrounding areas in this period (CPUESerr = 1.4 ±1.2 SE vs 0.6 ±1.1 SE ind.line-1.h-1, p<0.001; WPUESerr = 99.9 ±1.3 SE vs 33.7 ±1.2 SE g.line-1.h-1, p<0.001), and 17.3 and 23.3 times higher off-shore than on-shore (CPUESerr = 1.4 ±1.1 SE vs 0.1 ±1.2 SE ind.line-1.h-1, p<0.001; WPUESerr = 96.6 ±1.2 SE vs 4.1 ±1.2 SE g.line-1.h-1, p<0.001; S3 Table).

Between 2005 and 2014, Serranidae catch abundance and weight (Fig 4c, 4d) declined for off-shore fishermen, both in the reserve (-85% in CPUESerr from 3.2 to 0.5 ind.line-1.h-1, -78%Serr in WPUESerr from 226.5 to 49.0 g.line-1.h-1) as well as in surrounding areas (-81% in CPUESerr from 1.5 to 0.3 ind.line-1.h-1, -76%Serr in WPUESerr from 88.4 to 21.2 g.line-1.h-1; S4 Table). Anglers fishing on-shore outside the reserve also experienced declining CPUESerr (-86%, from 0.2 to 0.0 ind.line-1.h-1) and WPUESerr (-51%, from 6.6 to 3.2 g.line-1.h-1), whereas those fishing on-shore inside the reserve had increasing CPUESerr (+304%, from 0.0 to 0.1 ind.line-1.h-1) and WPUESerr (+17%, from 2.0 to 2.4 g.line-1.h-1) over time.

Labridae

Labridae represented 12% (700 ind.) of the total recorded catch, and 3% (25.2 kg) of the overall yield in weight. An average CPUELabr of 0.2 ±1.1 SE ind.line-1.h-1 and WPUESerr of 8.6 ±1.2 SE g.line-1.h-1 were recorded in 2005–2014 (considering all years). Labridae catch abundance and weight did not differ significantly between inside and outside the reserve in this period (CPUELabr = 0.2 ±1.3 SE vs 0.3 ±1.2 SE ind.line-1.h-1, p = 0.058; WPUELabr = 9.9 ±1.4 SE vs 7.9 ±1.3 SE g.line-1.h-1, p = 0.616), or among fishermen off-shore as compared with on-shore (CPUELabr = 0.3 ±1.2 SE vs 0.2 ±1.2 SE ind.line-1.h-1, p = 0.742; WPUELabr = 7.9 ±1.3 SE vs 9.4 ±1.4 SE g.line-1.h-1, p = 0.696; S3 Table).

Labridae catch abundance and weight declined in 2005–2014, both inside and outside of the reserve as well as for both on- and off-shore fishermen (Fig 4e and 4f; S4 Table). Along the shoreline, -65% in CPUELabr (from 0.3 to 0.1 ind.line-1.h-1) and -93% in WPUELabr (from 21.7 to 1.6 g.line-1.h-1) were estimated in the reserve, and -67% in CPUELabr (from 0.4 to 0.1 ind.line-1.h-1) and -80% in WPUELabr (from 12.5 to 2.5 g.line-1.h-1) in surrounding areas. For off-shore fishermen, -98% in CPUELabr (from 0.3 to 0.0 ind.line-1.h-1) and -83% in WPUELabr (from 9.5 to 1.6 g.line-1.h-1) were estimated within the reserve, and -70% in CPUELabr (from 0.5 to 0.2 ind.line-1.h-1) and -81% in WPUELabr (from 14.2 to 2.7 g.line-1.h-1) in surrounding waters.

Discussion

Effectiveness of the reserve

Understanding how marine species respond to conservation actions is crucial for successful management of fisheries resources. At the historical site of Cerbère-Banyuls, catch abundances and weights for recreational fishermen were respectively 40% and 50% higher within the buffer zone of partial protection in the reserve than in surrounding areas, indicating significant benefits of the reserve in supporting fishing yield. Fishing restrictions inside the reserve did not protect against the general pattern of decline in catch per unit effort (CPUE) observed in non-reserve areas in 2005–2014, but did support increasing weight per unit effort (WPUE) despite declining values outside the reserve. This indicates changing fishing yields in the reserve through time, with fewer catches overall but an increase in the size of fish that are caught. This finding differs from those reporting increases in catch abundance inside protected areas [12, 26, 35], which might be due to the relatively old age of the Cerbère-Banyuls reserve established in 1974. Indeed, marine reserves promote prolific marine populations, including large predatory species that take longer to re-establish and, through time, are expected to increasingly regulate the abundance of smaller assemblages via trophic cascades [17, 23, 25, 39–41]. For example, the Cerbère-Banyuls marine reserve hosts a recovering population of the large predator dusky grouper (Epinephelus marginatus) whose abundance has been increasing from 10 individuals in 1986 to more than 650 in 2020 (the population abundance over our study period being 202 in 2006 and 429 in 2014; www.gemlemerou.org). While the consequences of the loss of large predators for the dynamics of ecosystems is a global concern [42, 43], further investigation is necessary to evaluate the effects of the return of top predators on fisheries resources and marine biota at the Cerbère-Banyuls marine reserve.

Unequal benefits of the reserve

The benefits of the reserve in promoting fishing yields were limited to off-shore fishermen, whereas on-shore fishermen experienced on average ~4 times lower and declining fishing efficiencies, even within the perimeter of the reserve. While a spatial segregation of larger fish further from the shore could be anticipated, differing trajectories in yield over time indicated reserve benefits were restricted to off-shore fishermen (Fig 3). We did not test for differences in gear characteristics between fishermen (though our estimates of CPUE and WPUE accounted for differences in gear abundance). However, it is unlikely that a same group of users would use significantly different gear inside and outside of the reserve, and that this difference in gear effectiveness would be responsible for the growing yields recorded for off-shore fishermen in the reserve. Alternatively, differences in site accessibility, and therefore fishing pressure, may explain the restriction of reserve benefits to off-shore fishermen [11, 13, 44]. While off-shore fishing is restricted to boat users and segregates fishing effort in a two-dimensional space throughout the reserve, the near-shore is potentially accessible to all fishermen and concentrates fishing pressure on a few accessible sites (mostly along beeches and jetties) along the mono-dimensional stretch of the coastline (Fig 1). Moreover, near-shore habitats are typically more exposed to other forms of degradation that impact marine biota, including pollution and artificialization of the coastline [2, 6, 45]. Spearfishing, which is often pointed to as a major driver of fish decline in shallow water habitats [12, 25, 46], has been forbidden within the Cerbère-Banyuls marine reserve since 1974, and model simulations indicate that further reducing recreational fishing pressure by 50% could significantly improve stocks of targeted fish species in the area [47, 48]. In 2016, after the period covered by this study, additional restrictions were implemented to regulate recreational fishing pressure within the reserve, anglers being now limited to a maximum of 2 lines with up to 4 hooks if fishing on-shore and 8 hooks off-shore, and the total number of fishermen is now restricted to 1,000 free but mandatory annual permits. While the effects of these new measures on fishing efforts and yields remain to be evaluated, the number of fishing permits could further be reduced in the future as annual user-permit demands have been below the 1,000 threshold in recent years. Nevertheless, there is growing concern that declining fishing yields could jeopardize the popularity of recreational fishing, an emblematic activity in the region, with significant economic consequences for the associated sectors including bait markets, harbors, and tourism.

Changing catch composition

Spatial differences in fishing yield were found among dominant fish families, suggesting a spatial segregation of fish populations. Average catch abundance and weight in the 10-year period did not differ significantly between inside and outside of the reserve for Labridae and Sparidae, but were 2–3 times higher in the reserve for Serranidae. Similarly, equivalent levels of catch abundance and weight were found on- and off-shore for Labridae, while yields were twice higher off-shore for Sparidae and ~20 times higher for Serranidae. The heterogeneity of benthic habitats was previously found to influence spatial variability in fish assemblage abundance and composition more strongly than protection status at the Cerbère-Banyuls marine reserve [49]. However, contrasting trajectories in fishing yield among fish families over the study period indicated differences in exploitation and/or replenishment of populations and, therefore, changing fish assemblages. For Sparidae, fishing yields in weight increased over time in the reserve as well as in surrounding off-shore areas, while no benefits of the reserve were detected on catch abundances, indicating increasing harvested fish size through time. In contrast, catch abundance and weight declined in all areas for Labridae and Serranidae, except on-shore in the reserve where Serranidae catch abundance quadrupled over a decade.

The Sparidae family comprises several large and mobile species that are highly targeted by recreational and commercial fishermen [3, 12, 25, 27, 44, 46], including Dentex dentex, Sparus aurata, Lithognathus mormyrus, Pagrus pagrus, and Diplodus sargus (S1 Table). As such, the increasing yields recorded for Sparidae indicate the reserve was effective in supporting catches of large individuals from species of high-value to fisheries within the protected area as well as an apparent spillover benefit to adjacent off-shore areas as expected for effective marine reserves [23, 24, 35, 40]. In contrast, the species of Labridae and Serranidae found in the Cerbère-Banyuls marine reserve (S1 Table) are comparatively of low interest to fishermen, which might explain the small differences in yields found between reserve and non-reserve areas [23, 25, 40, 50]. The increasing catch abundance recorded for Serranidae along the shoreline may reflect the capacity of these relatively small, substrate-associated fishes to colonize habitats unoccupied by other species, notably large predatory fish from the family Sparidae [25]. Overall, our decadal-scale evaluation of recreational fishing yields at the Cerbère-Banyuls marine reserve indicates a progressive transfer in catch biomass in space from on-shore to off-shore, as well as in composition from smaller, less-targeted fish to larger species that are of higher value to fisheries.

Implications for management

Our study shows that surveys of recreational fishing activities can constitute effective alternatives for estimating fishery indicators (e.g. CPUE) compared with using data from commercial fisheries which often have higher uncertainties in declarations on fishing efforts and yields [8, 51, 52]. Our interactions with recreational fishermen also helped create dialogue between scientists, managers, and citizens, building awareness of local management actions to support participation and trust. Our results indicate that, 40 years after its establishment, fishing yields at the Cerbère-Banyuls marine reserve were still changing, implying complex ecological processes that establish on multi-decadal timescales following the creation of a reserve. Over the last decade, changes included shifting catch composition from smaller and less-targeted fish from families Serranidae and Labridae, which with decreasing yields appear in this context as ecological losers among local species, to larger Sparidae fishes that are of high value to fisheries and, with increasing yields, stand as ecological winners. The benefits of the reserve for local fisheries were however mostly restricted to fishermen fishing off-shore who, with increasing yields, stand as social winners of the current management plan, whereas fishermen on-shore, the social losers, suffered ~4 times lower and declining fishing yields. In addition, large confidence intervals surrounding estimates of fishing yields over recent years indicate that recorded increases in yields for off-shore fishermen may not be equally shared among users (Figs 2–4).

With mean CPUE ranging 0.4–4.0 ind.line-1.h-1 and mean WPUE ranging 30.5–543.4 g.line-1.h-1 (S4 Table), recreational fishing yields at the Cerbère-Banyuls marine reserve are within the range of those reported along the north-western Mediterranean coast [9]. Similarly, the decline in shallow-water fishing yield observed locally reflects the broader pattern of declining near-shore yields at the scale of the entire Mediterranean [1-3]. As such, local management outcomes at Cerbère-Banyuls can help define regional plans, though identifying how local success at the small scale of the reserve (650 ha) can be expanded to the broader scale of the marine park (4,010 km2) or that of the entire Gulf of Lion (Fig 1) remains a challenge. The possibility of multiplying the number of natural reserves like Cerbère-Banyuls to amplify marine protected area benefits and counter declining fishing yields and coastal degradation in the region is presently under discussion [53, 54]. Nevertheless, while the increasing yields for Sparidae attest for the positive effects of the reserve on stocks of targeted fish species, the decline of near-shore yields poses several challenges, including that of preserving the socio-economic benefits of fishery activities (both recreational and commercial) and maintaining access to resources for different user groups (both on- and off-shore), for which group-specific regulations could be enforced [3, 10, 11, 39]. Our findings indicate that current management plans do not benefit on-shore fishing, undermining equity in this emblematic activity that is historically accessible to all and particularly popular among vacationers, low-income, and retired people. Given increasing pressure on common-pool natural resources and growing socio-economic inequalities, this emerging issue needs to be prioritized in sustainable management actions [19, 22, 55].

List of the species recorded for each of the three major fish families caught by recreational fishermen around the Cerbère-Banyuls natural marine reserve.

(PDF)

Click here for additional data file.

Deviance table of generalized linear models characterizing changes in catch per unit effort and weight per unit effort of on-shore and off-shore recreational fishing, inside versus outside the Cerbère-Banyuls marine reserve.

(PDF)

Click here for additional data file.

Deviance table of generalized linear models comparing average catch per unit effort and weight per unit effort of on-shore and off-shore recreational fishing, inside versus outside the Cerbère-Banyuls marine reserve over the entire study period.

(PDF)

Click here for additional data file.

Yields in catch per unit effort and weight per unit effort estimated for on-shore and off-shore recreational fishing, inside and outside of the Cerbère-Banyuls marine reserve at the beginning and end of the survey.

(PDF)

Click here for additional data file.

Parameter estimates of generalized linear models characterizing trends in catch per unit effort and weight per unit effort of on-shore and off-shore recreational fishing, inside versus outside the Cerbère-Banyuls marine reserve.

(PDF)

Click here for additional data file.

Trends in catch per unit effort and weight per unit effort of recreational fishing inside versus outside the Cerbère-Banyuls marine reserve with unit effort expressed in line.hours and in hook.hours.

(PDF)

Click here for additional data file.

30 Sep 2020

PONE-D-20-23582

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

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Reviewer #1: Partly

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2. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: I Don't Know

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Reviewer #1: No

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Reviewer #1: Yes

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5. Review Comments to the Author

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Reviewer #1: This paper uses surveys of recreational anglers over a period of ten years to derive several indicators of fishing yield (Catch per unit effort (CPUE) and weight per unit effort (WPUE), in various zones in and around a marine reserve. The objective is to determine whether the reserve is resulting in improved fish stock status and recreational fishing yields, and to whom these benefits are distributed. They found that the average CPUE and WPUE per year were higher inside the reserve, but that over time, CPUE has declined both inside, and outside the reserve. However, they did find that WPUE is increasing inside the reserve, but declining outside. The increase in WPUE is only being experienced by offshore fishers in boats however, while the onshore anglers continue to experience a decline in WPUE, regardless of whether they are fishing inside or outside the reserve. This data has the potential to be a valuable source of evidence for decision making and the introduction and conclusion are well written, however the paper is missing some important details about the modelling methods and survey data, which make it difficult to assess the validity of the results obtained.

Main comments

My main concern is around the level of detail provided in the methods, particularly around the modelling. The methods state that the models are used to compare CPUE/WPUE in different zones and for species, however a lack of detail in the methods section means it is unclear how this was achieved, making it difficult to determine how robust the reported results are. More specifically:

The objective of the modelling is unclear: while the methods state the objective is to identify differences between different catch conditions, the results (eg Fig 1) show the models have been used to model trajectories of CPUE/WPUE for different groups, which were then compared (somehow) to ascertain their difference.

More information is needed on how many models were fitted (is there a separate model for each zone? taxonomic grouping?) and with what response and explanatory variables. For example:

- Is the response variable the CPUE/WPUE, subset by onshore/offshore etc, or is it the difference in CPUE/WPUE between onshore and offshore fishers?

- What are the explanatory variables considered, and retained in the final selected model? Are these the variables listed as ‘factors’ in Table S2?

Some details are provided in Table S2, but the caption, and particularly column names, are not informative. These issues could be relatively easily addressed by:

1. Explicitly stating the model’s purpose and response variables in the methods.

2. Explaining the model selection process undertaken in the methods

3. Including a tables of candidate, and selected, models in the supporting information to replace existing Table S2, along with specifically labelled explanatory and response variables (final models could even go in a table in the results depending on how many there are). (For examples of how to address points 2 and 3, you could refer to the main text and supporting information in Curnick DJ, Collen B, Koldewey HJ, Jones KE, Kemp KM, Ferretti F. 2020. Interactions between a large marine protected area, pelagic tuna and associated fisheries. Frontiers in Marine Science 7:318. )

4. Include some references to the literature around the methods used (general GLM literature as well as their specific use in fisheries for estimating indicators)

Estimation of CPUE and comparison of groups

Throughout the results, the authors compare average CPUE and WPUE between zones (onshore/offshore, in reserve/outside reserve), and provide a p value as evidence of statistical significance (eg line 239, 262 – 265 etc). It is not clear exactly how this p value was obtained, and they do not match p values reported in Table S4.

Please include a few sentences in the methods re. how the authors went from the models, to estimating the trajectories, and what test they used to compare average values to obtain the p values reported in the results.

Survey data

WPUE (Fig. 2 – 4 panel b) exhibits a marked expansion in confidence intervals post 2009. This pattern - combined with this result being somewhat unexpected (lines 351 – 352), and also informing some of the key arguments in the discussion - warrants some further attention in the discussion, along with some more information about the raw survey data (cleaning processes, figures showing the distribution – e.g. boxplots in the supporting information or as points in Figures 2 – 5)

Minor comments

Reporting of indicator values

The paper reports several different indicators across different zones (eg inshore/offshore, in reserve/outside reserve): Average catch abundance per year during the survey period, Average catch weight per year during the survey period (eg line 239 CPUE = 2.1 + 1.1 SE ind.line-1.h-1), CPUE trajectory over time, WPUE trajectory over time (eg +131%, from 222.3 to 514.0 g.line-1.h-1)

Because the authors are comparing the metrics over both space and time, it needs to be explicit which numbers refer to which indicator and type of comparison. Currently several different numbers are reported for CPUE but it is difficult to ascertain what the difference between these metrics are (some numbers refer to spatial averages, and others refer to trends over time). For example, rather than “fishing yield … was 1.4 times higher in terms of catch abundance (CPUE=2.1 ±1.1 SE vs 1.5 ±1.1 SE ind.line-1.h-1, p=0.002)” (lines 238 – 239), it would be clearer as something like “During the survey period, average catch abundance per year … was 1.4 times higher inside the reserve than outside (Ave. CPUE/year = 2.1 ±1.1 SE …)”.

Likewise, when comparing trajectories, explain that the decline is over time (rather than through space). This applies throughout the results section.

Lines 45 – 46 & 367 – 391: Discussion of unequal benefits. Could differences in gear (size/weight/type, rather than number of lines/hooks), or distribution of larger fish be another explanation for the differences in WPUE achieved by onshore/offshore fishers? If this is the case, further regulation of the reserve would be unlikely to achieve equitable distribution of benefits, so it may be important to cover this off before recommending regulation.

Lines 35 & 242: Begin by saying CPUE showed a ‘similar pattern’ – I think the objective was to explain that the CPUE trajectory is similar in both areas (in and out reserve), but comes across as saying the CPUE is similar to the results reported in the previous sentence. This gets very confusing as the previous sentence was reporting a positive effect. Suggest altering to something along the lines of ‘CPUE showed a declining trend over time, both inside and outside the reserve (reported figures here)’.

Lines 77 – 81: This paragraph provides important rationale for the research, but the logic could be articulated more explicitly – why do uncertainties about benefits pose regulatory challenges? How does identifying social and ecological winners address this challenge? How does it contribute to adaptive management? In addition, the discussion doesn’t address adaptive management specifically.

Lines 131 – 134: Description of the reserve’s value includes some unnecessary superlatives (e.g. exemplary, exceptional) which affect the objective tone used elsewhere.

Lines 189 – 191: The authors have made several assumptions about potential flaws in collecting survey data (1 – that respondees were honest, and 2 – that proportion of unreported catch remains consistent over time). These may be accepted assumptions for this method, but could be better supported with evidence/references and warrant a mention in the discussion.

Line 221: Variable ‘Time’ needs a unit – e.g. year, month, day?

Line 343 – 346 & 32: “At the historical site of Cerbère-Banyuls, catch abundances and weights for recreational fishermen were respectively 40% and 50% higher within the buffer zone of partial protection in the reserve than in surrounding areas, indicating significant benefits of the reserve in supporting fishing yield”. How is ‘significant benefit ’defined? Authors should rather discuss this in the context of the subsequent metrics, which indicate that while the overall CPUE may be higher inside the reserve, it is still declining steadily over time, so the benefits are not unequivocal. E.g., discuss the reserve benefits, with caveats, after reporting the other metrics as well.

Line 429 – 432: states that “ Our study shows that surveys of recreational fishing activities can constitute robust alternatives for estimating fishery indicators (e.g. CPUE) compared with using data from commercial fisheries …” , This statement should be rephrased to more accurately represent what was achieved. As no comparison is made with either commercial data or fisheries independent data, it is not possible to assess robustness, but the paper does demonstrate the use of a worthwhile alternate approach.

Lines 441 – 442: The terms social and ecological winners appear somewhat out of the blue here (although social and ecological protagonists are mentioned in the introduction). Defining these concepts and their implications in the introduction would strengthen this conclusion when it appears.

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12 Nov 2020

Editor’s comments:

In your revision please address carefully the comments/suggestions made by reviewer #1 regarding the background information on the modelling as well as the presentation and interpretation of the results.

Additional information is now provided on the modelling methods, results, and interpretation (see response to reviewer comments below).

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We note that one or more of the authors are employed by a commercial company: SEANEO.

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Thank you for these considerations about potential competing interests. In fact, contributing author N.S. participated in several of the survey campaigns of this study while under affiliations 1 & 2 (host laboratory at the University of Perpignan). We therefore amended her affiliations on the manuscript, indicating her previous affiliations with the host laboratory, and mentioning the private company SEANEO as her present address. Our competing interest statement therefore remains unchanged.

We also updated the affiliations for the three coauthors A.L.-D., J.D., and M.J. who were in similar cases (contributions to study under affiliations 1 & 2, with new current addresses).

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The maps in Fig 1 were produced using the open source program QGIS. This information is now provided in the figure caption (l. 103).

(1) Thank you for including your ethics statement on the online submission form:

This study involved interviews with recreational fishermen. The interviews were

conducted anonymously after informed consent for study participation from each

subject. The survey methodology and material was approved by the University of

Perpignan.

To help ensure that the wording of your manuscript is suitable for publication, would you please also add this statement at the beginning of the Methods section of your manuscript file.

This statement has been added at the beginning of the Methods section (l. 120-123).

(2) Thank you for providing additional information regarding the authors' affiliations. Could you please clarify whether author NS was affiliated with SEANO at the time of study?

Contributing author N.S. participated to this study while under affiliations 1 & 2 only (host laboratory at the University of Perpignan), with no affiliation with the commercial company SEANEO at the time of the study.

Reviewers' comments:

Reviewer #1: This paper uses surveys of recreational anglers over a period of ten years to derive several indicators of fishing yield (Catch per unit effort (CPUE) and weight per unit effort (WPUE), in various zones in and around a marine reserve. The objective is to determine whether the reserve is resulting in improved fish stock status and recreational fishing yields, and to whom these benefits are distributed. They found that the average CPUE and WPUE per year were higher inside the reserve, but that over time, CPUE has declined both inside, and outside the reserve. However, they did find that WPUE is increasing inside the reserve, but declining outside. The increase in WPUE is only being experienced by offshore fishers in boats however, while the onshore anglers continue to experience a decline in WPUE, regardless of whether they are fishing inside or outside the reserve. This data has the potential to be a valuable source of evidence for decision making and the introduction and conclusion are well written, however the paper is missing some important details about the modelling methods and survey data, which make it difficult to assess the validity of the results obtained.

Thank you for your in depth evaluation of our manuscript, and pointing out these issues. We have now revised the descriptions of our statistical analyses, clarified our overall study approach, and amended our interpretation of the results in the light of the comments (as detailed below).

Main comments

My main concern is around the level of detail provided in the methods, particularly around the modelling. The methods state that the models are used to compare CPUE/WPUE in different zones and for species, however a lack of detail in the methods section means it is unclear how this was achieved, making it difficult to determine how robust the reported results are. More specifically:

The objective of the modelling is unclear: while the methods state the objective is to identify differences between different catch conditions, the results (eg Fig 1) show the models have been used to model trajectories of CPUE/WPUE for different groups, which were then compared (somehow) to ascertain their difference.

More information is needed on how many models were fitted (is there a separate model for each zone? taxonomic grouping?) and with what response and explanatory variables. For example:

- Is the response variable the CPUE/WPUE, subset by onshore/offshore etc, or is it the difference in CPUE/WPUE between onshore and offshore fishers?

- What are the explanatory variables considered, and retained in the final selected model? Are these the variables listed as ‘factors’ in Table S2?

Some details are provided in Table S2, but the caption, and particularly column names, are not informative. These issues could be relatively easily addressed by:

1. Explicitly stating the model’s purpose and response variables in the methods.

A set of GLMs were indeed used to estimate trajectories of the response variables CPUE and WPUE inside versus outside of the reserve, and to identify temporal changes in yield for fishermen groups (on- and off-shore) and fish-families (Sparidae, Serranidae, Labridae). This general purpose is stated l. 212-215.

We initially implemented three-factor models (CPUE ~ Time × Reserve × Fishermen, and WPUE ~ Time × Reserve × Fishermen) to test for differences in reserve effects on trajectories among fishermen groups. However, lack of convergence indicated over-parametrization of some of the models, which we therefore split into simpler two-factor models (Time × Reserve) applied separately to on- and off-shore fishermen data). This is now specified l. 215-222.

For consistency, we report two-factor models (Time × Reserve, now better detailed in S2 Table) characterizing CPUE and WPUE trajectories for all species and fishermen (Fig. 2, model set 1 in S2 Table), and separately per fishermen group (Fig. 3, model sets 2-3 in S2 Table) and per fish family (Fig. 4, model sets 4-9 in S2 Table). We now also specify in the text that similar results were found when trajectories were estimated using two- and three-factor models, only the latter being reported in the manuscript (l. 222-225).

2. Explaining the model selection process undertaken in the methods

3. Including a tables of candidate, and selected, models in the supporting information to replace existing Table S2, along with specifically labelled explanatory and response variables (final models could even go in a table in the results depending on how many there are). (For examples of how to address points 2 and 3, you could refer to the main text and supporting information in Curnick DJ, Collen B, Koldewey HJ, Jones KE, Kemp KM, Ferretti F. 2020. Interactions between a large marine protected area, pelagic tuna and associated fisheries. Frontiers in Marine Science 7:318. )

As described above, we have now amended the manuscript text and S2 Table to clarify our statistical approach and design of the GLMs characterizing fishing yield trajectories. This includes a clear identification of explanatory and response variables (l. 215-222).

We did not perform additional model selection as used in GAMM modeling, though we now cite the suggested study by Curnick et al. in relation to the effects of the reserve for higher trophic level species (citation #41, manuscript l. 363).

4. Include some references to the literature around the methods used (general GLM literature as well as their specific use in fisheries for estimating indicators)

In addition to the already cited work by Ripley et al. 2019 on the MASS package from which the GLM function was used (citation #38 l. 233), we have now added reference to the book by Dunn & Smyth (2018) that provides numerous examples of GLM applications (citation #36 l. 212). We also added reference to work by Harrison et al. (2018) discussing the mentioned over-parametrization issue in GLMs (citation #37, l. 221).

Estimation of CPUE and comparison of groups

Throughout the results, the authors compare average CPUE and WPUE between zones (onshore/offshore, in reserve/outside reserve), and provide a p value as evidence of statistical significance (eg line 239, 262 – 265 etc). It is not clear exactly how this p value was obtained, and they do not match p values reported in Table S4.

Please include a few sentences in the methods re. how the authors went from the models, to estimating the trajectories, and what test they used to compare average values to obtain the p values reported in the results.

Average CPUE and WPUE over the entire study period (pooled over 2005-2014) were compared using a separate set of GLMs (different from those used to estimate trajectories). We previously did not detail this to avoid overloading of the manuscript, but we now include information on these tests in the Methods section l. 228-230 as well as in our new S3 Table.

Survey data

WPUE (Fig. 2 – 4 panel b) exhibits a marked expansion in confidence intervals post 2009. This pattern - combined with this result being somewhat unexpected (lines 351 – 352), and also informing some of the key arguments in the discussion - warrants some further attention in the discussion, along with some more information about the raw survey data (cleaning processes, figures showing the distribution – e.g. boxplots in the supporting information or as points in Figures 2 – 5)

Raw data on fishing yields are illustrated in S1 Fig. Yield data were characterized by high dispersion, which was taken into account in our analyses with the use of a GLM function for negative-binomial distribution (mentioned l. 231-233).

We preferred presenting average curves ±confidence-intervals in our figures in the manuscript, rather than average curves with raw data, given the difficulty to read the latter graphs due to broad axis value ranges (as illustrated in S1 Fig).

However, the remark of the reviewer on the increase in confidence interval in time is indeed interesting, as it indicates further dispersion of the data with years, or in other words, increasing disparity in yield among individual observations. This indicates the identified average increases in yields may not be equally distributed among off-shore fishermen, and is now mentioned in Discussion l. 452-454.

Minor comments

Reporting of indicator values

The paper reports several different indicators across different zones (eg inshore/offshore, in reserve/outside reserve): Average catch abundance per year during the survey period, Average catch weight per year during the survey period (eg line 239 CPUE = 2.1 + 1.1 SE ind.line-1.h-1), CPUE trajectory over time, WPUE trajectory over time (eg +131%, from 222.3 to 514.0 g.line-1.h-1)

Because the authors are comparing the metrics over both space and time, it needs to be explicit which numbers refer to which indicator and type of comparison. Currently several different numbers are reported for CPUE but it is difficult to ascertain what the difference between these metrics are (some numbers refer to spatial averages, and others refer to trends over time). For example, rather than “fishing yield … was 1.4 times higher in terms of catch abundance (CPUE=2.1 ±1.1 SE vs 1.5 ±1.1 SE ind.line-1.h-1, p=0.002)” (lines 238 – 239), it would be clearer as something like “During the survey period, average catch abundance per year … was 1.4 times higher inside the reserve than outside (Ave. CPUE/year = 2.1 ±1.1 SE …)”.

Likewise, when comparing trajectories, explain that the decline is over time (rather than through space). This applies throughout the results section.

Thank you. We added further clarifying terms throughout the Results section to account for this comment (l. 244, 248, 285, 315, 333).

Lines 45 – 46 & 367 – 391: Discussion of unequal benefits. Could differences in gear (size/weight/type, rather than number of lines/hooks), or distribution of larger fish be another explanation for the differences in WPUE achieved by onshore/offshore fishers? If this is the case, further regulation of the reserve would be unlikely to achieve equitable distribution of benefits, so it may be important to cover this off before recommending regulation.

While a segregation of larger fish further from the shore could be anticipated (that is, larger average WPUE off-shore), our estimations of fishing yield trajectories over time indicated the reserve benefits (increasing yields) were restricted to off-shore fishermen (now stated l. 375-377).

We did not test for differences in gear characteristics between fishermen (our estimates of CPUE and WPUE already accounting for differences in gear abundance). However, it is unlikely that a same group of users would use significantly different gear when fishing inside versus outside of the reserve, and that this difference in gear effectiveness would be responsible for the growing yields recorded for off-shore fishermen inside the reserve (now stated l. 377-382).

Lines 35 & 242: Begin by saying CPUE showed a ‘similar pattern’ – I think the objective was to explain that the CPUE trajectory is similar in both areas (in and out reserve), but comes across as saying the CPUE is similar to the results reported in the previous sentence. This gets very confusing as the previous sentence was reporting a positive effect. Suggest altering to something along the lines of ‘CPUE showed a declining trend over time, both inside and outside the reserve (reported figures here)’.

This has been corrected (l. 36 and 247-252).

Lines 77 – 81: This paragraph provides important rationale for the research, but the logic could be articulated more explicitly – why do uncertainties about benefits pose regulatory challenges?

The sentence now specifies in terms of implementing appropriate measures for resource durability and equitable access (l. 73-74).

How does identifying social and ecological winners address this challenge? How does it contribute to adaptive management? In addition, the discussion doesn’t address adaptive management specifically.

We have now changed this sentence to “… can refine regulatory strategies and help define win-win sustainable management for people and ecosystems” (l. 74-76).

Lines 131 – 134: Description of the reserve’s value includes some unnecessary superlatives (e.g. exemplary, exceptional) which affect the objective tone used elsewhere.

These superlatives have been removed.

Lines 189 – 191: The authors have made several assumptions about potential flaws in collecting survey data (1 – that respondees were honest, and 2 – that proportion of unreported catch remains consistent over time). These may be accepted assumptions for this method, but could be better supported with evidence/references and warrant a mention in the discussion.

There are no studies evaluating unreported catch of recreational fishermen in our study area. Similarly, there were no reasons to postulate that unreported catch would change over time over our study period. It is our preference to report these assumptions at once in the Methods section, and focus the Discussion on the findings of the study.

Line 221: Variable ‘Time’ needs a unit – e.g. year, month, day?

We now provide a range in years in the sentence: 2005-2014.

Line 343 – 346 & 32: “At the historical site of Cerbère-Banyuls, catch abundances and weights for recreational fishermen were respectively 40% and 50% higher within the buffer zone of partial protection in the reserve than in surrounding areas, indicating significant benefits of the reserve in supporting fishing yield”. How is ‘significant benefit ’defined? Authors should rather discuss this in the context of the subsequent metrics, which indicate that while the overall CPUE may be higher inside the reserve, it is still declining steadily over time, so the benefits are not unequivocal. E.g., discuss the reserve benefits, with caveats, after reporting the other metrics as well.

Given the multiple aspects of our study on reserve benefits for fish and fishermen groups, we opted for a hierarchical narrative in the discussion: starting from the simple effect of the reserve alone averaged over the 10 year period (l. 351-354), and unfolding with the sequential mention of differences in trajectories among fishing areas (l. 354-357), fishermen groups (l. 373-375) and fish families (l. 405-410).

Overall, it is hard to conclude if the reserve was not beneficial because catch numbers were declining, or if it was, because catch weights were increasing. Nevertheless, our study shows that while the two processes were taking place in the reserve, the benefits differed among fishermen and fish families, which needs to be taken into consideration in future management plans.

Line 429 – 432: states that “ Our study shows that surveys of recreational fishing activities can constitute robust alternatives for estimating fishery indicators (e.g. CPUE) compared with using data from commercial fisheries …” , This statement should be rephrased to more accurately represent what was achieved. As no comparison is made with either commercial data or fisheries independent data, it is not possible to assess robustness, but the paper does demonstrate the use of a worthwhile alternate approach.

The term “robust” has been replaced by “effective”.

Lines 441 – 442: The terms social and ecological winners appear somewhat out of the blue here (although social and ecological protagonists are mentioned in the introduction). Defining these concepts and their implications in the introduction would strengthen this conclusion when it appears.

The notion of winners/losers is now introduced in the introduction section l. 74-76.

17 Nov 2020

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

PONE-D-20-23582R1

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19 Nov 2020

PONE-D-20-23582R1

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

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