INTRODUCTIONTop

The European lobster, Homarus gammarus (Linnaeus), is a decapod crustacean species of great ecological and economic importance found in the northeastern Atlantic. Its distribution ranges from northern Norway to Morocco, but it is not present in the Baltic Sea (Holthuis 1991Holthuis L.B. 1991. Marine Lobsters of the world. An Annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Species Catalog. FAO Fish. Synop. 13: 1-292.). It also inhabits the Mediterranean and the western Black Sea, including the Istanbul Strait (Gönülal 2015Gönülal O. 2015. Spiny lobster (Palinurus elephas Fabricius, 1787) and common lobster (Homarus gammarus Linnaeus, 1758) fishing in the Agean Sea. Turkish Marine Research Foundation (TÜDAV).).

The species is listed in the Mediterranean in Appendix 3 (Protected fauna species) of the Berne convention on the Conservation of European Wildlife and Natural Habitats (e.g. Lloret and Riera 2008Lloret J., Riera V. 2008. Evolution of a Mediterranean coastal zone: human impacts on the marine environment of Cape Creus. Environ. Manage. 42: 977-988.). This benthic animal is a nocturnal and territorial species found mainly on rocky substrates which feeds on a wide range of prey such as blue mussels, hermit crabs and Polychaeta (Bertran and Le Calvez 1988Bertran R., Le Calvez J.C. 1988. Contenus stomacaux dejeunes homards européens capturés en pêche à pied à Blainville (Ouest-Cotentin). In: International Council for the Exploration of the Sea Council Meeting. 1-12 pp., Prodöhl et al. 2006Prodöhl P.A., Jørstad K.E., Triantafyllidis A., et al. 2006. European lobster-Homarus gammarus. In: Genetic Impact of Aquaculture Activities on Native Populations. Norwegian Institute of Marine Research. Final Sci. Rep. pp. 91-98.). Individuals can weigh more than 7 kg and the largest males and females may live up to 42 and 72 years, respectively (Sheehy et al. 1999Sheehy M.R.J., Bannister R.C.A., Wickins J.F., et al. 1999. New perspectives on the growth and longevity of the European lobster (Homarus gammarus). Can. J. Fish. Aquat. Sci. 56: 1904-1915.). H. gammarus and the American lobster H. americanus (Milne Edwards), found on the northwest coast of the Atlantic, are the only two species representing their genus within the family Nephropidae, and are anatomically very similar (Wahle et al. 2013Wahle R.A., Castro K.M., Tully O., et al. 2013. Homarus. In: Phillips B. (ed.), Lobsters: biology, management, aquaculture and fisheries, 2nd ed. Wiley, New York, pp. 221-258.).

With 5194 t of total landings in 2014 (FAO 2017FAO. 2017. FISHSTAT J: FAO fishery and aquaculture global statistics.), the European lobster is a significant fishery resource throughout its range. It is an important source of income and employment for artisanal and coastal fisheries considering its high commercial value. It is targeted by trap fisheries in northern Europe, mainly in the United Kingdom, Ireland, and northern France (Bennett and Lovewell 1977Bennett D.B., Lovewell S.R.J. 1977. The Effects of Pot Immersion Time on Catches of Lobsters Homarus gammarus (L) in the Welsh Coast Fishery. Great Britain Ministry of Agriculture, Fisheries and Food., Bennet et al. 1993Bennet D., Casey J., Dare P. et al. 1993. Identification biogéographique des principaux stocks exploités en Manche, relations avec ceux des régions voisines., Browne et al. 2001Browne R.M., Mercer J.P., Duncan M.J. 2001. An historical overview of the Republic of Ireland’s lobster (Homarus gammarus Linnaeus) fishery, with reference to European and North American (Homarus americanus Milne Edwards) lobster landings. In: Coastal Shellfish-A Sustainable Resource. Springer, 49-62 pp.). The traps have various designs, shapes, and sizes, and they can capture singly (one buoy for one trap) or in a gang (several buoys for several tens of traps) (Cobb and Castro 2006Cobb J.S., Castro K.M. 2006. Homarus species. In: Philips B.F. (ed.), Lobsters: biology, management, aquaculture, and fisheries, Blackwell Publishing Ltd., Oxford, UK. pp. 310-339.). Trap fishing is primarily performed inshore, often less than 5 km from the coast.

Genetic analyses using different molecular markers (mitochondrial DNA and allozymes) were conducted to look for genetic differentiation across the entire range of H. gammarus (Jørstad et al. 2005Jørstad K.E., Farestveit E., Kelly E., et al. 2005. Allozyme variation in European lobster (Homarus gammarus) throughout its distribution range. N. Z. J. Mar. Freshw. Res. 39: 515-526., Triantafyllidis et al. 2005Triantafyllidis A., Apostolidis A.P., Katsares V., et al. 2005. Mitochondrial DNA variation in the European lobster (Homarus gammarus) throughout the range. Mar. Biol. 146: 223-235.). They indicated the presence of genetic subpopulations, broadly defined as Mediterranean, Atlantic, northern Norway, and the Netherlands (Triantafyllidis et al. 2005Triantafyllidis A., Apostolidis A.P., Katsares V., et al. 2005. Mitochondrial DNA variation in the European lobster (Homarus gammarus) throughout the range. Mar. Biol. 146: 223-235.). This information, coupled with studies showing limited movement of adult lobsters (Moland et al. 2011Moland E., Olsen E.M., Knutsen H., et al. 2011. Activity patterns of wild European lobster Homarus gammarus in coastal marine reserves: implications for future reserve design. Mar. Ecol. Prog. Ser. 429: 197-207.), a short larval stage (Bennet et al. 1993Bennet D., Casey J., Dare P. et al. 1993. Identification biogéographique des principaux stocks exploités en Manche, relations avec ceux des régions voisines.), and possibly small-scale larval dispersal (Schmalenbach and Buchholz 2010Schmalenbach I., Buchholz F. 2010. Vertical positioning and swimming performance of lobster larvae (Homarus gammarus) in an artificial water column at Helgoland, North Sea. Mar. Biol. Res. 6: 89-99.), suggests that the Mediterranean subpopulation of the European lobster population should be considered separately.

Artisanal fisheries or small-scale coastal fisheries are particularly important in the Mediterranean (Farrugio et al. 1993Farrugio H., Oliver P., Biagi F. 1993. An overview of the history, knowledge, recent and future research trends in Mediterranean fisheries. Sci. Mar. 57: 105-119., Guyader et al. 2013Guyader O, Berthou P, Koutsikopoulos C., et al. 2013. Small scale fisheries in Europe: A comparative analysis based on a selection of case studies. Fish. Res. 140: 1-13.), where they constitute about 80% of the fishing fleet (European Commission 2004European Commission. 2004. Mediterranean: guaranteeing sustainable fisheries. Fishing in Europe 21: 1-12.). Fishermen operate with small boats near the coast (0-200 m depth) and adapt their practices to resource availability: a wide variety of gear is used, targeting a diversity of species that change in space and time (Colloca et al. 2004Colloca F., Crespi V., Cerasi S., et al. 2004. Structure and evolution of the artisanal fishery in a southern Italian coastal area. Fish. Res. 69: 359-369., Leleu et al. 2014Leleu K., Pelletier D., Charbonnel E., et al. 2014. Métiers, effort and catches of a Mediterranean small-scale coastal fishery: The case of the Côte Bleue Marine Park. Fish. Res. 154: 93-101.). Along the Mediterranean coast, the European lobster is not a target species and is more often a by-catch that occurs in trammel nets targeting the common spiny lobster Palinurus elephas (Fabricius) (Marin 1987Marin J. 1987. Exploitation, biologie et dynamique du stock de langouste rouge de Corse, Palinurus elephas Fabricius. Ph. D. thesis. Université d’Aix-Marseille II., Quetglas et al. 2004Quetglas A., Gaamour A., Reñones O., et al. 2004. Common spiny lobster (Palinurus elephas Fabricius 1787) fisheries in the western Mediterranean: A comparison of Spanish and Tunisian fisheries. Boll. Soc. Hist. Nat. Balears 47: 63-80.) or in gillnets targeting fish (Gönülal 2015Gönülal O. 2015. Spiny lobster (Palinurus elephas Fabricius, 1787) and common lobster (Homarus gammarus Linnaeus, 1758) fishing in the Agean Sea. Turkish Marine Research Foundation (TÜDAV).). Spiny lobster fishing in the Mediterranean accounts for a large part of the income generated during the fishing season, but it is showing increasing signs of overexploitation (Goñi and Latrouite 2005Goñi R., Latrouite D. 2005. Review of the biology, ecology and fisheries of Palinurus spp. species of European waters: Palinurus elephas (Fabricius, 1787) and Palinurus mauritanicus (Gruvel, 1911). Cah. Biol. Mar. 46: 127-142., Pere 2012Pere A. 2012. Déclin des populations de langouste rouge et baisse de la ressource halieutique en Corse: causes et perspectives. Ph. D. thesis, Université de Corse, 478 pp.). Given the equivalent high commercial value of H. gammarus, this species is of potential interest for fishermen wishing to diversify crustacean-fishing activities in the Mediterranean. In Corsica (NW Mediterranean), the overall fleet can be described as artisanal, which makes it fully representative of Mediterranean small-scale fisheries (Marengo et al. 2016Marengo M., Pere A., Marchand B., et al. 2016. Catch variation and demographic structure of common dentex (Sparidae) exploited by Mediterranean artisanal fisheries. Bull. Mar. Sci. 92: 191-206.).

While the literature on the ecology and fishing of H. gammarus in the Atlantic is abundant, no single study provides comprehensive information regarding its exploitation in the Mediterranean Sea and its potential for the development of specific fisheries. The aim of this study was therefore to obtain a better understanding of the exploitation pattern and to collect initial biological information on the European lobster population around Corsica. More specifically, we analysed (1) the catch per unit of effort (CPUE), including the dynamics of the catches in space, time, and water depth; (2) the total annual production of the island; (3) the distribution; and (4) the potential abundance of the species.

MATERIALS AND METHODSTop

Study area

The study was conducted in Corsica, France (northwestern Mediterranean, geographical sub-area 8, 42°N and 9°E, Fig. 1). It covers 1047 km of coastline and is characterized by high environmental heterogeneity with the occurrence of the seagrass Posidonia oceanica (Linnaeus) Delile, meadows, rocky bottoms and sandy shores. Geomorphology and bathymetric range are highly heterogeneous, with a narrow continental shelf, submarine canyons and deep water along the western coast, as well as wide expanses of shallow waters on the eastern coast, along the Tyrrhenian Sea (Pluquet 2006Pluquet F. 2006. Evolution récente et sédimentation des plates-formes continentales de la Corse. Ph. D. thesis, Université de Corse Pascal Paoli, 300 pp.).

Fishery data

Data were collected by scientific observers on board fishing vessels for eight consecutive years (2006 to 2013) during the main fishing period (April to September). Around the island, fishing boats are widely dispersed over 34 harbours and smaller sheltered areas where boats can be anchored during the fishing season. In the first step, the fishermen of the “active” boats were identified, and the data were updated each year (Pere 2012Pere A. 2012. Déclin des populations de langouste rouge et baisse de la ressource halieutique en Corse: causes et perspectives. Ph. D. thesis, Université de Corse, 478 pp.). “Active” boats are those which have a real activity among all the boats having a fishing licence.

A total of 922 fishing trips were performed during the eight-year sampling period (Table 1). Fishing activity is practised near the coast and average time at sea is 6-7 hours (Pere et al. 2008Pere A., Lejeune P., Pelaprat C. 2008. Suivi scientifique de la pêche langoustière Corse. Rapport final-Année 2007. Off. Environ. Corse, Stareso Fr.). Each year, the daily observed fishing trips were chosen at random by a network of four to five scientific observers spread all over the island. During the monitoring period, 30% of the active fleet was sampled at least once every year and 75% of the active fleet was sampled during the whole study period. A total of 4523 hauls were sampled during the study, corresponding to 2379 km of net. All the nets were observed during a sampled fishing trip. Data related to technical characteristics (length, mesh size, type of net and mesh composition) and utilization (location, depth, soak time) were collected. After capture, each European lobster was measured (carapace length, CL) to the nearest mm, and sex was recorded. As it was impossible to precisely weigh individuals on board commercial vessels, weight was estimated from available length-weight relationships (Latrouite 2001Latrouite D. 2001. Le homard (Homarus gammarus) du nord golfe de Gascogne-Manche ouest (divisions VIIe + VIIIa du CIEM). Contrat Ifremer/MAPA – n° 99-I1-03-01. 13 pp.) for males and females.

Scientific survey data

The independent fishery survey was performed in 2013 and 2014 in the west of the Cap Corse area with a 12-m professional fishing boat using selective inkwell pot, “Conquetois” type European lobster traps (height 43 cm; length 65 cm; width 50 cm; 20 cm diameter opening on the top). The survey area included a fishing no-take area (Cantonnement de Nonza, Fig. 1). Traps were arranged in lines of three spaced 20 m apart. Salted sardine was used as a bait in a specific pocket inside the trap. The spatial sampling scheme was randomly stratified, covering various substrata and a bathymetry ranging between 30 and 120 m depth. For each trip 20 lines were deployed, totalling 180 lines and 540 traps over the survey period. The sampling area covered fishing grounds located northwest of the Island used by fishermen from Centuri harbour, and the Nonza no-take area was sampled at its northern and southern parts for operational reasons. Soak time was fixed to four days in order to maximize the catch rate (Smith and Tremblay 2003Smith S.J., Tremblay M.J. 2003. Fishery-independent trap surveys of lobsters (Homarus americanus): design considerations. Fish. Res. 62: 65-75.). Captured lobsters were measured for CL and total length to the nearest mm, weighed (g), sexed, then tagged using T-bar tags and released at the capture point.

Data analysis

Catches (CPUE) were expressed in density (number) and biomass (weight) of European lobsters per 50 m of net. All nets were taken into account (trammel nets, gillnets and combined nets), and CPUE included commercial lobsters, small lobsters that returned alive to the sea, and dead lobsters (Table 2). For the spatial analyses, the island was divided into six fishing areas (Fig. 1), designated as Ajaccio, Calvi, Cap Corse, Plaine, Porto, and Bonifacio (Pere et al. 2008Pere A., Lejeune P., Pelaprat C. 2008. Suivi scientifique de la pêche langoustière Corse. Rapport final-Année 2007. Off. Environ. Corse, Stareso Fr.). These fishing areas were determined based on geomorphologic criteria (e.g. differences in depth and habitat type).

The combination of continuous and categorical variables in generalized linear models (GLM) allows the relative influence of variables affecting catch rates to be evaluated (Maynou et al. 2003Maynou F., Demestre M., Sánchez P. 2003. Analysis of catch per unit effort by multivariate analysis and generalised linear models for deep-water crustacean fisheries off Barcelona (NW Mediterranean). Fish. Res. 65: 257-269.). The CPUE data were log transformed [log(CPUE + 1)] prior to all the analyses to meet the normality requirements of the test, and a Gaussian distribution was used (Santos et al. 2014Santos M.N., Coelho R., Lino P.G. 2014. Standardized CPUE for swordfish (Xiphias gladius) caught by the Portuguese pelagic longline fishery in the North Atlantic. Collect Vol Sci Pap ICCAT 70: 1783-1791.). Adjusted sums of squares (Type III) were used to calculate the F-value and p-values. The main factor effects and interactions were determined using the GLM (Minitab 2000Minitab I.N.C. 2000. MINITAB statistical software. Minitab Release 13.). The explanatory variables considered and tested with the GLM were years (2009-2013, 8 levels), months (April–September, 6 levels), depths (0 to >100 m depth, 6 levels) and fishing areas (Ajaccio, Bonifacio, Calvi, Cap Corse, Plaine, Porto, 6 levels). Within the GLM, all possible pairs of interactions were tested. For example, the two-way and three-way interactions between these factors were investigated using a “forward selection” procedure.

Specific comparisons between the factors were performed using post hoc Tukey honest significance difference (HSD) tests. Finally, catch and effort spatial distributions were mapped using a geographical information system (QGIS 2.6.1). Fishing effort was estimated as the annual number of fishing trips and mean length of net hauled per fishing trip. The annual total yield estimate was derived from the catch data and the fishing effort calculated above (for more details, see Pere 2012Pere A. 2012. Déclin des populations de langouste rouge et baisse de la ressource halieutique en Corse: causes et perspectives. Ph. D. thesis, Université de Corse, 478 pp.). The positionS of the trap lines and lobster catches were represented on the map of Corsica and bathymetry was added using ArcGIS 10.2 (Fig. 1B).

RESULTSTop

Catch variation

According to the variance analysis produced by the GLM, fishing areas, depths and months were the factors that had a significant effect on the catch rates of H. gammarus (Table 3), but CPUE values were not significantly related to the factor years. Interactions between months and depths as well as between years, months and depths were significant. Statistically significant interactions (p<0.01) between months and fishing areas were also detected. Tukey’s HSD test revealed that CPUE differed significantly across fishing areas, being much higher at Bonifacio than at Ajaccio, Calvi, Cap Corse and Plaine (Fig. 2A).

Catches of this species showed a seasonal peak in April and July (Fig. 2B) and a wide range of bathymetric distribution with values ranging from 10 (minimum) to 164 m depth (maximum). The maximum exploitation depth was between 40 and 60 m (Fig. 2C). CPUE differed significantly among depths (p<0.05), and was smaller at 0-20 m, 20-40 m and >100 than at 40-60 m (Fig. 2C).

Spatial distribution of catches

A heterogeneous distribution of catches of the European lobster was observed around Corsica. The highest CPUEs were mainly located in the south of Corsica (Bonifacio) (Fig. 3), while the lowest were found in the east of Corsica (Plaine) and Cap Corse.

Total catches

An annual fluctuation of catches was observed. During the study period, the estimated mean annual total production was 6.7 t (±1.6 t se). Over time, the time-series analysis highlighted several trends. Recent years have been marked by strong intra- and inter-annual fluctuations in catches and two main trends were observed: (i) a relatively stable yield from 2006 to 2009 and (ii) an increase in 2011, with maximum production that reached 9.8 t/year (Fig. 4). The peak was mainly due to catches in a favourable fishing area (Bonifacio). After 2011, fluctuations occurred and catches dropped to 5 to 7 t per year (2012-2013).

Size distribution

Between 2006 and 2013, a total of 244 specimens were sampled. The size of H. gammarus ranged from 61 to 200 mm CL (Fig. 5). Three predominant size classes were detected: 100-120, 120-140 and 140-160 (Fig. 5).

Fishery independent survey

The fishery independent survey showed an extremely low catch rate over the study period, with only one lobster caught out of the 540 deployed traps, resulting in a mean catch rate of 0.002±0.043 lobsters per trap. This individual was caught in May 2013 at 35 m depth (Fig 1B). It was a male of 150 mm CL, 383 mm total length and 2.5 kg.

DISCUSSIONTop

Marin (1987)Marin J. 1987. Exploitation, biologie et dynamique du stock de langouste rouge de Corse, Palinurus elephas Fabricius. Ph. D. thesis. Université d’Aix-Marseille II. estimated European lobster and European spider crab Maja squinado (Herbst) production at 5 to 10 t per year in Corsica. This result is consistent with average annual catches estimated at 6.7 t in this study. Based on FAO data for the period (2006-2013), production in France was estimated at 550 t per year (Atlantic and Mediterranean coasts, FAO 2017FAO. 2017. FISHSTAT J: FAO fishery and aquaculture global statistics.), so Corsica represents about 1.2% of total captures of this species at the national level. The European lobster is an accessory catch in Corsica, but it can constitute a significant part of landings, especially in the south of the island. Production on the French Mediterranean coast is estimated at 5.5 t, confirming the general underestimates of FAO data in the Mediterranean in general and also in Corsica (Le Manach et al. 2011Le Manach F., Dura D., Pere A., et al. 2011. Preliminary estimate of total marine fisheries catches in Corsica, France (1950-2008. In: Harper S., Zeller D. (eds), Fisheries catch reconstructions: Islands, Part II. Fish. Cent. Res. Rep. 19. Univ. British Columbia.).

Historical records of the Mediterranean show catastrophic declines in the landings of clawed lobster in the 20th century (Dow et al. 1980Dow R.L., Cobb J.S., Phillips B.F. 1980. The clawed lobster fisheries. Biol. Manag. Lobsters 2: 265-316.). According to Cobb and Castro (2006)Cobb J.S., Castro K.M. 2006. Homarus species. In: Philips B.F. (ed.), Lobsters: biology, management, aquaculture, and fisheries, Blackwell Publishing Ltd., Oxford, UK. pp. 310-339., this trend persists today. Statistics of this type should be considered with caution, since it is difficult to evaluate the landings of artisanal small-scale fisheries and they are often underestimated. However, it has been clearly demonstrated that specific lobster stocks in the Mediterranean have suffered from overexploitation, with significant declines in catches, e.g. in Spain (Lloret and Riera 2008Lloret J., Riera V. 2008. Evolution of a Mediterranean coastal zone: human impacts on the marine environment of Cape Creus. Environ. Manage. 42: 977-988.) and in the Adriatic (Lotze et al. 2011Lotze H.K., Coll M., Dunne J.A. 2011. Historical changes in marine resources, food-web structure and ecosystem functioning in the Adriatic Sea. Mediterr. Ecosyst. 14: 198-222.).

In addition to overexploitation, a pattern of low abundance is evident throughout the Mediterranean compared with Atlantic fisheries. Temperature changes might play a major role for the H. gammarus population in its southern distribution, with excessively high temperatures leading to reduced population abundance (Triantafyllidis et al. 2005Triantafyllidis A., Apostolidis A.P., Katsares V., et al. 2005. Mitochondrial DNA variation in the European lobster (Homarus gammarus) throughout the range. Mar. Biol. 146: 223-235.). High temperature could especially affect fragile, newly settled European lobsters. Although specific observation of the early benthic phase of the European lobster in the wild is still lacking (Linnane et al. 2001Linnane A., Ball B., Mercer J.P., et al. 2001. Searching for the early benthic phase (EBP) of the European lobster: a trans-European study of cobble fauna. Hydrobiologia 465: 63-72.), H. gammarus post larvae are assumed to settle and remain cryptic in shelter-providing rocky substrata and to emerge from their shelters only once they have reached between 25 and 40 mm CL (Linnane et al. 2000aLinnane A., Ball B., Munday B., et al. 2000a. On the occurrence of juvenile lobster Homarus gammarus in intertidal habitat. J. Mar. Biol. Assoc. U.K. 80: 375-376.,bLinnane A., Mazzoni D., Mercer J.P. 2000b. A long-term mesocosm study on the settlement and survival of juvenile European lobster Homarus gammarus L. in four natural substrata. J. Exp. Mar. Biol. Ecol. 249: 51-64., Ball et al. 2001Ball B., Linnane A., Munday B., et al. 2001. The effect of cover on in situ predation in early benthic phase European lobster Homarus gammarus. J. Mar. Biol. Assoc. U.K. 81: 639-642.). Hence, high temperatures in shallow coastal waters during the summer period in the Mediterranean could play a major role in juvenile survival and recruitment success. Moreover, the European lobster may be sensitive to climate change, as rising temperature is the most important factor driving shifts in its range of distribution, and increased abundance is found at higher latitudes and decreased abundance at lower latitudes (Caputi et al. 2013Caputi N., Lestang S., Frusher S., et al. 2013. The impact of climate change on exploited lobster stocks. In: Philips B.F. (ed.), Lobsters: biology, management, aquaculture, and fisheries, Blackwell Publishing Ltd., Oxford, UK, pp. 84-112., Green et al. 2014Green B.S., Gardner C., Hochmuth J.D., et al. 2014. Environmental effects on fished lobsters and crabs. Rev. Fish Biol. Fish. 24: 613-638.).

Catchability of the European lobster is affected by numerous factors, including feeding behaviour and moulting status (Wahle et al. 2013Wahle R.A., Castro K.M., Tully O., et al. 2013. Homarus. In: Phillips B. (ed.), Lobsters: biology, management, aquaculture and fisheries, 2nd ed. Wiley, New York, pp. 221-258.). Thus, high catches in April could be the consequence of an increase in lobster feeding activity with increasing temperature (Bennett and Lovewell 1977Bennett D.B., Lovewell S.R.J. 1977. The Effects of Pot Immersion Time on Catches of Lobsters Homarus gammarus (L) in the Welsh Coast Fishery. Great Britain Ministry of Agriculture, Fisheries and Food., Lizárraga-Cubedo et al. 2015Lizárraga-Cubedo H.A., Tuck I., Bailey N., et al. 2015. Scottish lobster fisheries and environmental variability. ICES J. Mar. Sci. 72: i211-i224.). The decline in catch rates from May to June may be attributable to moulting activity (Miller 1990Miller R.J. 1990. Effectiveness of crab and lobster traps. Can. J. Fish. Aquat. Sci. 47: 1228-1251., Sheehy et al. 1999Sheehy M.R.J., Bannister R.C.A., Wickins J.F., et al. 1999. New perspectives on the growth and longevity of the European lobster (Homarus gammarus). Can. J. Fish. Aquat. Sci. 56: 1904-1915.), and the following peak in CPUE in July is consistent with the increased feeding activity observed during the post-moult period.

The size structures of Corsican catches (range 61-200 mm CL; mean size 127 mm CL) seem to shift towards larger individuals compared with those coming from east Britain (range 78-115 mm CL; mean size 94 mm CL), South Wales (range 90-146 mm CL; mean size 109 mm CL), Scotland (range 62-140 mm CL; modal size 95 mm CL), and Brittany (range 60-160 mm CL) (Howard 1987Howard A.E. 1987. Estimates of lobster (Homarus gammarus) fecundity from East and West Britain. ICES CM., Lizárraga-Cubedo et al. 2003Lizárraga-Cubedo H.A., Tuck I., Bailey N.P et al. 2003. Comparisons of size at maturity and fecundity of two Scottish populations of the European lobster, Homarus gammarus. Fish. Res. 65: 137-152., Laurans et al. 2009Laurans M., Fifas S., Demaneche S., et al. 2009. Modelling seasonal and annual variation in size at functional maturity in the European lobster (Homarus gammarus) from self-sampling data. ICES J. Mar. Sci. 66: 1892-1898.). The minimum conservation reference size (MCRS), which is usually broadly equivalent to the mean size at first maturity, was imposed under EU fisheries legislation at 87 mm CL for the Atlantic (EC Reg., 850/98) and 105 mm CL for the Mediterranean (EC Reg., 1967/2006). In this study, 80% of the lobsters sampled were above the MCRS, indicating that the individuals were mostly mature.

No individual smaller than 60 mm CL was caught, and similar observations were made by Latrouite (2001)Latrouite D. 2001. Le homard (Homarus gammarus) du nord golfe de Gascogne-Manche ouest (divisions VIIe + VIIIa du CIEM). Contrat Ifremer/MAPA – n° 99-I1-03-01. 13 pp. and Prodöhl et al. (2006)Prodöhl P.A., Jørstad K.E., Triantafyllidis A., et al. 2006. European lobster-Homarus gammarus. In: Genetic Impact of Aquaculture Activities on Native Populations. Norwegian Institute of Marine Research. Final Sci. Rep. pp. 91-98. in the Atlantic. This result could be linked to gear selectivity, because many European lobsters in the Mediterranean are caught in trammel nets with a large-mesh size (62-83 mm) for the inner panel (Campillo and Amadei 1978Campillo A., Amadei J. 1978. Premieres donnees biologiques sur la langouste de Corse, Palinurus elephas Fabricius. Rev. Trav. Inst. Pêch. Marit. 42: 347-373.). Finally, the cryptic behaviour and limited movement of juveniles could better explain the absence of small and young European lobsters.

In the Atlantic, H. gammarus primarily occurs from low intertidal zones to depths usually no greater than 50 m (Holthuis 1991Holthuis L.B. 1991. Marine Lobsters of the world. An Annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Species Catalog. FAO Fish. Synop. 13: 1-292.). For example, the mean water depth of the lobster catches was 37.5 m on the Basque continental shelf (Galparsoro et al. 2009Galparsoro I., Borja Á., Bald J., et al. 2009. Predicting suitable habitat for the European lobster (Homarus gammarus), on the Basque continental shelf (Bay of Biscay), using Ecological-Niche Factor Analysis. Ecol. Model. 220: 556-567.). In Corsica, bathymetric analysis revealed that the highest abundance was between 40 and 80 m. This deeper distribution is potentially due to the high surface temperature in the Mediterranean Sea, which can affect this cold-water species.

In this study, 65% of the animals were encountered in “spiny lobster nets”, which are gears defined by the target species P. elephas and by their use (soak time ≥2 d and depth ≥50 m) (Pere 2012Pere A. 2012. Déclin des populations de langouste rouge et baisse de la ressource halieutique en Corse: causes et perspectives. Ph. D. thesis, Université de Corse, 478 pp.). Other Mediterranean small-scale fisheries display the same trend. In Languedoc-Roussillon (northwestern Mediterranean, France), H. gammarus is mainly found in gillnets for various finfish and in trammel nets for crustaceans (Guillou et al. 2002Guillou A., Lespagnol P., Ruchon F. 2002. La pêche aux petits métiers en Languedoc-Roussillon en 2000-2001. Ifremer. Convention de participation au programme PESCA (PIC) DIRAM - IFREMER n° 00/3210040/YF, et Convention de recherche Région Languedoc-Roussillon - IFREMER n° 00/1210041/YF). Around the Tabarca Marine Reserve (southwestern Mediterranean, Spain), H. gammarus and Scorpaena scrofa Linnaeus are accessory species caught in trammel nets for Palinurus (mean water depth=64 m; mean soak time=46 h) (Forcada et al. 2009Forcada A., Valle C., Sánchez-Lizaso J.L., et al. 2009. Structure and spatio-temporal dynamics of artisanal fisheries around a Mediterranean marine protected area. ICES J. Mar. Sci. 67: 191-203.).

The bathymetric distribution of European lobsters and the nature of the fishing net in which it is caught provide information on adult habitats, which essentially consist of rocky and coralligenous substrates where natural protective structures (micro-caves, holes and crevices) are numerous. It has been reported that the most suitable habitats for the European lobster are locations at the boundary between sedimentary and rocky bottoms (Galparsoro et al. 2009Galparsoro I., Borja Á., Bald J., et al. 2009. Predicting suitable habitat for the European lobster (Homarus gammarus), on the Basque continental shelf (Bay of Biscay), using Ecological-Niche Factor Analysis. Ecol. Model. 220: 556-567.). In Corsica, these kinds of substrates are widely distributed around the island (except on the east side, Martin et al. 2014Martin C.S., Giannoulaki M., De Leo F., et al. 2014. Coralligenous and maërl habitats: predictive modelling to identify their spatial distributions across the Mediterranean Sea. Sci. Rep. 4: 50-73.) and are particularly common in the Strait of Bonifacio, where the continental shelf has the particularity of being very large (Buron et al. 2012Buron K., Monville I., Jousseaume M., et al. 2012. Inventaires biologiques et analyse écologique des habitats marins patrimoniaux. Sites Natura 2000 en mer du lot Corse Extrême Sud. Volet III : Sites Natura 2000 FR9402015 - Bouches de Bonifacio, Iles des Moines; FR9400587 - Iles Cerbicale et franges littorale et FR9400591 - Plateau de Pertusato/Bonifacio et Iles Lavezzi. Rapport EVEMar-Stareso-Sintinelle/Agence des Aires Marines Protégées. 261 pp.).

The spatial approach identified an area of high abundance in the south, precisely in the Strait of Bonifacio. In the south of the island, potentially suitable habitats for adults coupled with (i) a considerable larval pool from the Bonifacio Strait Natural Reserve (Santoni et al. 2008Santoni M.C., Negre N., Culioli J.M. 2008. Expérimentation de la pêche à la nasse à l’intérieure et en périphérie du cantonnement de pêche de Bonifacio. Etude réalisée en convention avec la prud’homie de Bonifacio. Réserve Naturelle des Bouches de Bonifacio - Département Parc Marin Internationnal de l’Office de l’Environnement de la Corse. OEC Ed. 42 pp.), (ii) favourable early life history traits such as short pelagic larval duration (Bennet et al. 1993Bennet D., Casey J., Dare P. et al. 1993. Identification biogéographique des principaux stocks exploités en Manche, relations avec ceux des régions voisines., Goñi and Latrouite 2005Goñi R., Latrouite D. 2005. Review of the biology, ecology and fisheries of Palinurus spp. species of European waters: Palinurus elephas (Fabricius, 1787) and Palinurus mauritanicus (Gruvel, 1911). Cah. Biol. Mar. 46: 127-142., Øresland and Ulmestrand 2013Øresland V., Ulmestrand M. 2013. European lobster subpopulations from limited adult movements and larval retention. ICES J. Mar. Sci. 70: 532-539.) and (iii) advantageous hydrological features such as eddies (Gérigny et al. 2015Gérigny O., Coudray S., Lapucci C., et al. 2015. Small-scale variability of the current in the Strait of Bonifacio. Ocean Dyn. 65: 1165-1182.) could explain the high abundance of the European lobster. For H. gammarus, larval self-recruitment mechanisms have already been discussed (Schmalenbach and Buchholz 2010Schmalenbach I., Buchholz F. 2010. Vertical positioning and swimming performance of lobster larvae (Homarus gammarus) in an artificial water column at Helgoland, North Sea. Mar. Biol. Res. 6: 89-99., Øresland and Ulmestrand 2013Øresland V., Ulmestrand M. 2013. European lobster subpopulations from limited adult movements and larval retention. ICES J. Mar. Sci. 70: 532-539.). Elsewhere, several spots around the island also show high European lobster abundance. Despite the lack of favourable habitats on the eastern coast, several wrecks known by fishermen provide shelters for lobsters (Reveche 1979Reveche C. 1979. La mise en place de la gestion rationnelle d’un stock exploité: l’exemple du comité local des pêches maritimes (CLPM) de Blainville sur Mer et des fonds à Homard de la Côte Ouest du Cotentin. In: Journées d’étude Aquaculture Extensive et Repeuplement,-Brest, 29-31 Mai 1979.). The north of the island is marked by low yields. This result is somewhat surprising because the northern grounds are potentially suitable for the European lobster, with the occurrence of rocky bottoms and coralligenous assemblages (Pluquet 2006Pluquet F. 2006. Evolution récente et sédimentation des plates-formes continentales de la Corse. Ph. D. thesis, Université de Corse Pascal Paoli, 300 pp., Bonacorsi 2012Bonacorsi M. 2012. Caractérisation des peuplements benthiques du Cap Corse. Ph. D. thesis. Université de Corse, France. 1-159 pp.). However, this low European lobster catch rate is corroborated by the fishery-independent data from the scientific survey performed in west Cap Corse. By contrast, spiny lobsters are abundant in these areas, which is consistent with their preference for such grounds (Marin 1987Marin J. 1987. Exploitation, biologie et dynamique du stock de langouste rouge de Corse, Palinurus elephas Fabricius. Ph. D. thesis. Université d’Aix-Marseille II., Pere 2012Pere A. 2012. Déclin des populations de langouste rouge et baisse de la ressource halieutique en Corse: causes et perspectives. Ph. D. thesis, Université de Corse, 478 pp.).

Various habitats and depths were covered seasonally by the spatially randomly stratified sampling scheme, but only one lobster was caught out of 540 traps. It is generally expected for experimental fishing to achieve lower catch rates that follow a randomly stratified sampling scheme than fishery-dependent data obtained by fishing effort concentrated on specific areas. Experimental Atlantic fishing surveys had a mean catch rate of between 0.8 and 3 lobsters per trap in Norway (Moland et al. 2013Moland E., Olsen E.M., Knutsen H., et al. 2013. Lobster and cod benefit from small-scale northern marine protected areas: inference from an empirical before–after control-impact study. Proc. R. Soc. B 280: 20122679.) and between 0.29 and 1.32 lobster per trap in England (Addison 1995Addison J.T. 1995. Influence of behavioural interactions on lobster distribution and abundance as inferred from pot-caught samples. In: ICES Marine Science Symposia. International Council for the Exploration of the Sea, Copenhagen, Denmark, pp. 294-300.). A relatively low abundance of 0.09 lobsters per trap was observed in the south of the Bay of Biscay (Galparsoro et al. 2009Galparsoro I., Borja Á., Bald J., et al. 2009. Predicting suitable habitat for the European lobster (Homarus gammarus), on the Basque continental shelf (Bay of Biscay), using Ecological-Niche Factor Analysis. Ecol. Model. 220: 556-567.). The low abundance in west Cap Corse corroborates the low catch rates obtained with fishery-dependent data for the same area. Unfortunately, to our knowledge no single survey using specific lobster traps has been performed in the Mediterranean, thus making it impossible to make such a comparison.

Onboard monitoring of small-scale fisheries is an important tool that contributes to the suitable exploitation of various key species from the perspective of the island’s fishery management (Le Manach et al. 2011Le Manach F., Dura D., Pere A., et al. 2011. Preliminary estimate of total marine fisheries catches in Corsica, France (1950-2008. In: Harper S., Zeller D. (eds), Fisheries catch reconstructions: Islands, Part II. Fish. Cent. Res. Rep. 19. Univ. British Columbia., Marengo et al. 2016Marengo M., Pere A., Marchand B., et al. 2016. Catch variation and demographic structure of common dentex (Sparidae) exploited by Mediterranean artisanal fisheries. Bull. Mar. Sci. 92: 191-206.). Despite the reporting of greater landings in Fishstat statistics since 2006, the abundance of H. gammarus seems to remain low and stable (Wahle et al. 2013Wahle R.A., Castro K.M., Tully O., et al. 2013. Homarus. In: Phillips B. (ed.), Lobsters: biology, management, aquaculture and fisheries, 2nd ed. Wiley, New York, pp. 221-258.). In Corsica, the lack of historical landing datasets prevents us from concluding on the evolution of the stock and on the current population’s health status. However, the occurrence of this non-target species in selective (trap) and non-selective (trammel net) fishing gear around the island is low. In this context, we cannot reasonably expect that the targeting of European lobster would be an efficient solution for diversification of the artisanal fishery at the scale of the whole island.

ACKNOWLEDGEMENTSTop

The authors would like to thank A. Astrou for his involvement in building the database, L. Fullgrabe, who improved the language of the manuscript, and K. Buron, O. Gérigny and L.N. Michel for their scientific advice and suggestions, which also contributed to the improvement of the manuscript. We thank P. Sartor (Editorial Board) and two anonymous reviewers for their constructive comments on the previous version of the manuscript. The fishery-dependent surveys were funded by the Office de l’Environnement de la Corse and the Direction des Pêches Maritimes et de l’Aquaculture for 2006-2013, and the scientific fishery-independent survey was funded by FEDER funds for 2013-2015. The two fishery surveys would not have been carried out without the effective collaboration of the Corsican fishermen and the work of many scientific fishery observers.

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