Movements and spatial distribution of an endangered fish (Sciaena umbra) within a marine protected area.

The brown meagre (Sciaena umbra) is an endangered species, which requires specific protection measures to ensure its conservation. These measures need to be informed by high-quality scientific knowledge on their space use patterns. Here, we used acoustic telemetry to assess its seasonal movement patterns and habitat use within a marine protected area (MPA). Our results suggested that S. umbra is a highly sedentary species (home range < 1.0 km2) and, therefore, the MPA is extensive enough to protect the local population. Their population was discretely distributed in two main areas within the MPA, which was likely a result of habitat segregation and density-dependent movements. The temporal variability of their movements further uncovered when and where spawning occurs (mainly, but probably not only, in the fully protected area in June) and indicated that spillover of this species is limited but still possible. Overall, we highlight the importance of MPAs in the recovery of S. umbra, we advocate the need to perpetuate the current national fishing bans and extend it to other countries in the Mediterranean region, and we emphasize that considering the fine-scale movements of S. umbra in future management actions is key to achieving a successful recovery of their populations.

Harbours as unique environmental sites of multiple anthropogenic stressors on fish hormonal systems.

Fish development and acclimation to environmental conditions are strongly mediated by the hormonal endocrine system. In environments contaminated by anthropogenic stressors, hormonal pathway alterations can be detrimental for growth, survival, fitness, and at a larger scale for population maintenance. In the context of increasingly contaminated marine environments worldwide, numerous laboratory studies have confirmed the effect of one or a combination of pollutants on fish hormonal systems. However, this has not been confirmed in situ. In this review, we explore the body of knowledge related to the influence of anthropogenic stressors disrupting fish endocrine systems, recent advances (focusing on thyroid hormones and stress hormones such as cortisol), and potential research perspectives. Through this review, we highlight how harbours can be used as "in situ laboratories" given the variety of anthropogenic stressors (such as plastic, chemical, sound, light pollution, and invasive species) that can be simultaneously investigated in harbours over long periods of time.

Environmental DNA metabarcoding reveals and unpacks a biodiversity conservation paradox in Mediterranean marine reserves.

Although we are currently experiencing worldwide biodiversity loss, local species richness does not always decline under anthropogenic pressure. This conservation paradox may also apply in protected areas but has not yet received conclusive evidence in marine ecosystems. Here, we survey fish assemblages in six Mediterranean no-take reserves and their adjacent fishing grounds using environmental DNA (eDNA) while controlling for environmental conditions. We detect less fish species in marine reserves than in nearby fished areas. The paradoxical gradient in species richness is accompanied by a marked change in fish species composition under different managements. This dissimilarity is mainly driven by species that are often overlooked by classical visual surveys but detected with eDNA: cryptobenthic, pelagic, and rare fishes. These results do not negate the importance of reserves in protecting biodiversity but shed new light on how under-represented species groups can positively react to fishing pressure and how conservation efforts can shape regional biodiversity patterns.

Holistic investigation of shore angler profiles to support marine protected areas management.

Marine Recreational Fishing (MRF) is growing worldwide in scientific interest, as evidenced by the increasing number of dedicated publications. Studies on the impacts and benefits to socio-ecosystems and mental health are driving this gradual awareness. In the Mediterranean, MRF is currently responsible for 10% of the catches though, in the context of small-scale fisheries decline, it may become dominant. Sustaining this activity represents a universal challenge for the future of mankind. However, the potential influence of anglers' heterogeneity on both the environment and the Catch Per Unit of Effort (CPUE), used internationally to evaluate fish harvest and as a stocks indicator, hinders MRF management. In addition, little data is available on onshore fishing, while the number of practitioners may increase sharply in the context of a socio-economic crisis, especially in marine protected areas (MPA). We conducted a year-round survey in 2017-2018 in a French Mediterranean MPA subject to tourist flows, during which 144 onshore anglers were polled by semi-directive interviews. We used a typology based on technical and socio-economic data of anglers to describe their behaviour diversity and its influence on CPUE and, more broadly, the marine environment in multi-species fisheries. We characterised four onshore angler profiles, segregated in space and time, including one identified as at risk of causing impacts. Our analyses support using total and per species CPUE independently of profiles to evaluate onshore MRF harvest, except for four species. CPUE seem based on the profiles' skills and self-regulation which induce similar yields between anglers in the absence of control. This demonstrates the importance of taking into account angler behaviour, as each profile could react differently to management actions and highlights that CPUE could be used to detect the effect of changes in regulation in the framework of adaptive management. Our results support that MRF regulations should be simplified and homogenised at the national level and cooperation with fishing shops in MPA co-management should be promoted.

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.

Long-Distance Benefits of Marine Reserves: Myth or Reality?

Long-distance (>40-km) dispersal from marine reserves is poorly documented; yet, it can provide essential benefits such as seeding fished areas or connecting marine reserves into networks. From a meta-analysis, we suggest that the spatial scale of marine connectivity is underestimated due to the limited geographic extent of sampling designs. We also found that the largest marine reserves (>1000km2) are the most isolated. These findings have important implications for the assessment of evolutionary, ecological, and socio-economic long-distance benefits of marine reserves. We conclude that existing methods to infer dispersal should consider the up-to-date genomic advances and also expand the spatial scale of sampling designs. Incorporating long-distance connectivity in conservation planning will contribute to increase the benefits of marine reserve networks.

Spatial distribution of juvenile fish along an artificialized seascape, insights from common coastal species in the Northwestern Mediterranean Sea.

Along the littoral, a growing number of anthropogenic structures have caused substantial habitat destruction. Despite their detrimental impact, these constructions could play a role in the functioning of coastal ecosystems. The objective of this work was to assess the distribution of juvenile coastal fish along a seascape composed of various natural and artificial habitats in order to determine the potential role of coastal infrastructures as juvenile habitat. We surveyed juvenile populations on various infrastructures and natural sites along a 100 km shoreline of the French Mediterranean coast. Juvenile densities varied according to the level of artificialization of the sites. Densities were the highest on coastal defense structures, intermediate in natural sites and lowest in harbors. Focusing inside harbors revealed highly variable densities depending on the type of habitat, with densities on ripraps or jetties that were equivalent to those of natural sites. Our results underline the importance of anthropogenic structures as potential juvenile habitats, which is too often not considered in management plans.

Shallow rocky nursery habitat for fish: Spatial variability of juvenile fishes among this poorly protected essential habitat.

Coastal nursery habitats are essential for the renewal of adult fish populations. We quantified the availability of a coastal nursery habitat (shallow heterogeneous rocky bottoms) and the spatial variability of its juvenile fish populations along 250km of the Catalan coastline (France and Spain). Nurseries were present in 27% of the coastline, but only 2% of them benefited from strict protection status. For nine taxa characteristic of this habitat, total juvenile densities varied significantly between nursery sites along the coastline, with the highest densities being found on the northern sites. Recruitment level (i.e. a proxy of nursery value) was not explained by protection level, but it was moderately and positively correlated with an anthropization index. Patterns of spatial variations were taxa-specific. Exceptional observations of four juveniles of the protected grouper Epinephelus marginatus were recorded. Our data on habitat availability and recruitment levels provides important informations which help to focus MPA management efforts.

Spatial behavior of two coral reef fishes within a Caribbean marine protected area.

A better understanding of the key ecological processes of marine organisms is fundamental to improving design and effective implementation of marine protected areas (MPAs) and marine biodiversity. The movement behavior of coral reef fish is a complex mechanism that is highly linked to species life-history traits, predation risk and food resources. We used passive acoustic telemetry to study monthly, daily and hourly movement patterns and space use in two species, Schoolmaster snapper (Lutjanus apodus) and Stoplight parrotfish (Sparisoma viride). We investigated the spatial overlap between the two species and compared intra-specific spatial overlap between day and night. Presence-absence models showed different diel presence and habitat use patterns between the two species. We constructed a spatial network of the movement patterns, which showed that for both species when fish were detected by the array of receivers most movements were made around the coral reef habitat while occasionally moving to silt habitats. Our results show that most individuals made predictable daily crepuscular migrations between different locations and habitat types, although individual behavioral changes were observed for some individuals across time. Our study also highlights the necessity to consider multiple species during MPA implementation and to take into account the specific biological and ecological traits of each species. The low number of fish detected within the receiver array, as well as the intraspecific variability observed in this study, highlight the need to compare results across species and individuals to be used for MPA management.

Diel and seasonal movement pattern of the dusky grouper Epinephelus marginatus inside a marine reserve.

Temporal movement patterns and spawning behaviour of the dusky grouper Epinephelus marginatus were investigated using depth and temperature sensors combined to acoustic telemetry. Results showed that these fish are year-round resident, remaining inside the fully protected area of the marine reserve of Cerbère-Banyuls (65 ha) and display a diurnal activity pattern. Records from depth sensors revealed that groupers range inside small, distinct, and individual territories. Individual variations in habitat depth are only visible on a seasonal scale, i.e., between the spawning season and the rest of the year. In fact, during summer months when the seawater temperature exceeded 20 °C, tagged groupers made vertical spawning migrations of 4-8 m in amplitude. These vertical migrations are characteristic of the reproductive behaviour of dusky groupers, during which they release their gametes. The results are notable for the implementation of management rules in marine protected areas, such as reduced navigation speed, boating or attendance during spawning season.

Contrasting fish behavior in artificial seascapes with implications for resources conservation.

Artificial reefs are used by many fisheries managers as a tool to mitigate the impact of fisheries on coastal fish communities by providing new habitat for many exploited fish species. However, the comparison between the behavior of wild fish inhabiting either natural or artificial habitats has received less attention. Thus the spatio-temporal patterns of fish that establish their home range in one habitat or the other and their consequences of intra-population differentiation on life-history remain largely unexplored. We hypothesize that individuals with a preferred habitat (i.e. natural vs. artificial) can behave differently in terms of habitat use, with important consequences on population dynamics (e.g. life-history, mortality, and reproductive success). Therefore, using biotelemetry, 98 white seabream (Diplodus sargus) inhabiting either artificial or natural habitats were tagged and their behavior was monitored for up to eight months. Most white seabreams were highly resident either on natural or artificial reefs, with a preference for the shallow artificial reef subsets. Connectivity between artificial and natural reefs was limited for resident individuals due to great inter-habitat distances. The temporal behavioral patterns of white seabreams differed between artificial and natural reefs. Artificial-reef resident fish had a predominantly nocturnal diel pattern, whereas natural-reef resident fish showed a diurnal diel pattern. Differences in diel behavioral patterns of white seabream inhabiting artificial and natural reefs could be the expression of realized individual specialization resulting from differences in habitat configuration and resource availability between these two habitats. Artificial reefs have the potential to modify not only seascape connectivity but also the individual behavioral patterns of fishes. Future management plans of coastal areas and fisheries resources, including artificial reef implementation, should therefore consider the potential effect of habitat modification on fish behavior, which could have key implications on fish dynamics.

Combined effects of levels of protection and environmental variables at different spatial resolutions on fish assemblages in a marine protected area.

The links between species-environment relations and species' responses to protection are unclear, but the objectives of marine protected areas (MPAs) are most likely to be achieved when those relations are known and inform MPA design. The components of a species' habitat vary with the spatial resolution of the area considered. We characterized areas at two resolutions: 250 m(2) (transect) and approximately 30,000 m(2) (seascape). We considered three categories of environmental variables: substrate type, bottom complexity, and depth. We sought to determine at which resolution habitat characteristics were a better predictor of abundance and species composition of fishes and whether the relations with environmental variables at either resolution affected species' responses to protection. Habitat features accounted for a larger proportion of spatial variation in species composition and abundances than differences in protection status. This spatial variation was explained best by habitat characteristics at the seascape level than at the transect level. Species' responses to protected areas were specific to particular seascape characteristics, primarily depth, and bottom complexity. Our method may be useful for prioritizing marine areas for protection, designing MPAs, and monitoring their effectiveness. It identified areas that provided natural shelter, areas acting as buffer zones, and areas where fish species were most responsive to protection. The identification of such areas is necessary for cost-effective establishment and monitoring of MPAs.

Acoustic telemetry survey of the dusky grouper (Epinephelus marginatus) in the Marine Reserve of Cerbère-Banyuls: informations on the territoriality of this emblematic species.

The Marine Reserve of Cerbère-Banyuls represents a refuge for dusky grouper populations (Epinephelus marginatus). Density survey and observation of the reproductive behavior of this species have been conducted there for the last 30 years. However, information on the behavior or habitat utilization for this species during winter time is limited. We conducted acoustic telemetry surveys from September 2005 to June 2006 in the Marine Reserve of Cerbère-Banyuls for a dusky grouper population. Six groupers were tagged and followed within the integral reserve and in the adjacent partial reserve. The results from the tracking of these individuals indicated that the largest dusky groupers are sedentary fishes with a high degree of site fidelity and a high sensitivity to variations of environmental conditions such as storms. This together with the life history characteristics of this species encourages the conservation and the protection of essential habitats of this species. Acoustic tracking coupled to the use of a fixed Kernel can be proposed to follow fish populations.

Spatial assessment of fishing effort around European marine reserves: implications for successful fisheries management.

We examined the spatial dynamic of artisanal fishing fleets around five European marine protected areas (MPAs) to derive general implications for the evaluation of MPAs as fisheries management tools. The coastal MPAs studied were located off France, Malta and Spain and presented a variety of spatial designs and processes of establishment. We developed a standardized methodology to define factors influencing effort allocation and to produce fishing effort maps by merging GIS with geostatistical modelling techniques. Results revealed that in most cases the factors "distance to the no-take", "water depth", and "distance to the port" had a significant influence on effort allocation by the fishing fleets. Overall, we found local concentration of fishing effort around the MPA borders. Thus, neglecting the pattern of fishing effort distribution in evaluating MPA benefits, such as spillover of biomass, could hamper sound interpretation of MPAs as fisheries management tools.

Demographic and genetic structures of white sea bream populations (Diplodus sargus, Linnaeus, 1758) inside and outside a Mediterranean marine reserve.

We studied the white sea bream (Diplodus sargus), a protandrous hermaphroditic fish, in two protected and unprotected areas in southwestern France. We observed a significant difference in the demographic structure between the two areas. Females were present in two different age distributions inside and outside the marine reserve with younger females outside. This suggests plasticity in the age of sexual inversion in the case of an exploited population. Genetic differentiation was weak and apparent at only one locus of 26 surveyed (FST = 0.007, p = 0.04). Our data suggest that gene flow between the two areas is important, or the separation between the two sites is recent. Our data on the white sea bream show that fishes inside and outside the marine reserve are very similar genetically, which means that the 'reserve effect' is truly a demographic one, not the result of genetic differences.

[Heterozygosity and selective value: the case of the marine fish, Diplodus sargus (Linné, 1758)]. / Hétérozygotie et valeur sélective: le cas du poisson marin, Diplodus sargus (Linné, 1758).

A total of 842 white sea bream (Diplodus sargus), sampled in Banyuls-sur-Mer, were analysed to test 'genotype-phenotype' relationship for various characters related to the fitness. The results show significant differences (MLH and FIS) for the age according to the sex between females carrying out and not carrying out sexual inversion. This suggests an overdominance for old females and a genetic sex determination. The individuals laying very early during the period of reproduction are also differentiated from the individuals reproducing later in the season. These results suggest either a stable calendar of laying in time separating the individuals genetically reproducing precociously from the others and this by differential selection and/or genetic drift either a Wahlund effect among cohorts.