SARDINES EAST ATLANTIC (excluding the Mediterranean Sea) Sardina pilchardus. Sometimes known as European pilchard SUMMARY

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1 SARDINES EAST ATLANTIC (excluding the Mediterranean Sea) Sardina pilchardus Sometimes known as European pilchard SUMMARY Sardines are a pelagic, schooling fish (meaning they swim in groups near the surface) which grow fast and become sexually mature within two years. Sardines are an important commercial species in the eastern Atlantic, off the European and African coasts. Total annual catch is approximately 1 million tons, with many populations having low levels of abundance with some overexploited. Management measures are in place throughout their range, but it is unclear whether the current measures will meet sustainability goals. Most Sardines are caught using purse seines and pelagic trawls, which cause minimal habitat damage. The level of bycatch in these fisheries is not known, but is likely moderate. Criterion Points Final Score Color Life History Abundance Habitat Quality and Fishing Gear Impacts Management 2.00 Bycatch 2.00 Final Score 2.15 Color

2 LIFE HISTORY Core Points (only one selection allowed) If a value for intrinsic rate of increase ( r ) is known, assign the score below based on this value. If no r-value is available, assign the score below for the correct age at 50% maturity for females if specified, or for the correct value of growth rate ('k'). If no estimates of r, age at 50% maturity, or k are available, assign the score below based on maximum age Intrinsic rate of increase <0.05; OR age at 50% maturity >10 years; OR growth rate <0.15; OR maximum age >30 years Intrinsic rate of increase = ; OR age at 50% maturity = 5-10 years; OR a growth rate = ; OR maximum age = years Intrinsic rate of increase >0.16; OR age at 50% maturity = 1-5 years; OR growth rate >0.30; OR maximum age <11 years. Sardines grow quickly throughout their range in the eastern Atlantic Ocean, with growth rates, k, ranging from (Fishbase 2010). Growth in Sardine shows a latitudinal decline across the eastern Atlantic from the English Channel to north Morocco and growth in the Mediterranean tends to be lower than in other parts of the eastern Atlantic (Silva et al 2008). In the Iberian and Biscay Seas of the northeast Atlantic, the average growth rate, k, is 0.73 (Silva et al. 2008). In the Mediterranean, growth rates range from (Bigot and Roos 2009, Bellido et al. 2009, Quintanilla et al. 2009, Omar and Hachem 2009, Giannoulaki et al. 2009). Length at sexual maturity for Sardines ranges from cm (Fishbase 2010, Bigot and Ross 2009, Bellido et al. 2009, Quintanilla et al. 2009, Omar and Hachem 2009, Giannoulaki et al. 2009, Amenzoui et al. 2006) and they can reach a length of 25 cm (FAO 2010). Most Sardine mature by age 1 and all fish reach sexual maturity by age 2 (Bellido et al. 2009, Quintanilla et al. 2009, Giannoulaki et al.2009, Silva et al. 2006). Maximum ages of Sardine are variable, ranging from 4 years in the Aegean Sea to 14 years in the North Sea and English Channel (Fishbase 2010, Silva et al. 2008). In both the Mediterranean and Africa, few fish older than age 6 are caught (Bigot and Roos 2009, Bellido et al. 2009, Quintanilla et al. 2009, Santojanni and Cingolani 2009, FAO 2007). Overall, these life history characteristics indicate a score of 3. Points of Adjustment (multiple selections allowed) Species has special behaviors that make it especially vulnerable to fishing pressure (e.g., spawning aggregations; site fidelity; segregation by sex; migratory bottlenecks; unusual attraction to gear; etc.). Sardine is a schooling and migratory species (FAO 2010). Schools of juvenile fish tend to be found in inshore waters, often associated with estuaries or rivers (Tsagarakis K 2008).

3 There is also evidence that Sardines form short-lived spawning aggregations (Ganias 2008). These life history characteristics make Sardine particularly vulnerable to fishing pressure Species has a strategy for sexual development that makes it especially vulnerable to fishing pressure (e.g., age at 50% maturity >20 years; sequential hermaphrodites; extremely low fecundity) Species has a small or restricted range (e.g., endemism; numerous evolutionarily significant units; restricted to one coastline; e.g., American lobster; striped bass; endemic reef fishes). Sardines are found in the eastern Atlantic from the North Sea southward along the eastern coast of Europe, into the Mediterranean and Black Seas, and also along the north coast of Africa to Senegal (FAO 2010, Parrish et al ). The Sardine is a coastal, pelagic species found primarily from depths of 22 to 55 meters, but can also be found down to depths of 100 to 150 meters (FAO 2010, Parrish et al. 1989). Relative to other species, Sardine is considered to have a small range Species exhibits high natural population variability driven by broad-scale environmental change (e.g. El Nino; decadal oscillations). Spawning activity, growth, and recruitment of Sardines have all been linked to environmental influences (Ganias 2009, Catalan et al. 2006, Guisande 2001, Solaria et al. 2010). In the southwestern Adriatic Sea (Mediterranean), small pelagic species, including Sardines, have been shown to be affected by environmental variability (GFCM 2006). Variability of Sardine abundance has also been related to environmental factors in the North Atlantic region (Guisande et la. 2004, Alheit and Hagen 1997) Species does not have special behaviors that increase ease or population consequences of capture OR has special behaviors that make it less vulnerable to fishing pressure (e.g., species is widely dispersed during spawning) Species has a strategy for sexual development that makes it especially resilient to fishing pressure (e.g., age at 50% maturity <1 year; extremely high fecundity). In the eastern North Atlantic, Sardines spawn from mid March to November in the English Channel, with peaks in May/June and October, and throughout much of the year in the Bay of Biscay, with peaks from March to May (Stratoudakis et al. 2007). In the Iberian seas spawning extends from autumn to early spring (Stratoudakis et al. 2007). Spawning in Mediterranean waters occurs from September to May, peaking from November to February (Somarkasis et al. 2006, Palomera et al. 2007), and spawning off the African coast occurs from November to June (FAO 2010). Spawning takes place mainly in inshore waters over the continental shelf, primarily at temperatures of C (Palomera et al. 2007, Somarkasis et al. 2006). The duration of spawning, and therefore the number of batches laid by females, is dependent on environmental conditions and fish

4 size (Morello and Arneri 2009, Amenzoui et al. 2006). Sardines can spawn up to 15 times or batches per season (Morello and Arneri 2009). Mean batch fecundity ranges from 5,000 to 9,000 eggs in the Mediterranean (Somarkasis et al. 2006, Morello and Arneri 2009), and 14,000 to 34,000 eggs in the Atlantic (Somarkasis et al. 2006). Sardine is considered to be a moderately fecund species and therefore points are not added Species is distributed over a very wide range (e.g., throughout an entire hemisphere or ocean basin; e.g., swordfish; tuna; Patagonian toothfish) Species does not exhibit high natural population variability driven by broad-scale environmental change (e.g., El Nino; decadal oscillations) Points for Life History ABUNDANCE Core Points (only one selection allowed) Compared to natural or un-fished level, the species population is: 1.00 Low: Abundance or biomass is <75% of BMSY or similar proxy (e.g., spawning potential ratio). Sardines represent an important fishery in the eastern Atlantic (FAO 2010). From 1950s overall catches of Sardines initially increased, peaking in 1976 (1, 315, 685 tons) and again in 1990 (1, 525, 184 tons) (FAO 2010). Since 1990, however, Sardine catches have declined, with annual catches around 1,000,000 tons over the last decade (FAO 2010). In the European North Atlantic (Bay of Biscay and western Iberian Seas), spawning stock biomass has been variable but shows an overall decline since the early 1980s (ICES 2010a). Since 2006, spawning stock biomass has declined due to a lack of recruitment and is currently 33% below the long term average (ICES 2010a). Recruitment for 2008 and 2009 though are thought to be near the long-term average (ICES 2010a). Off Africa, abundance has been declining since 2000 for the northern population and currently biomass is at about 20% of BMSY. For the southern population, biomass has been increasing and is currently above BMSY. The northern fisheries are considered to be overexploited whereas the southern fishery is not fully exploited (FAO 2007). Given that the majority of Sardine populations in the eastern Atlantic are considered to have low abundance levels, a score of 1 was awarded.

5 2.00 Medium: Abundance or biomass is % of BMSY or similar proxy; OR population is approaching or recovering from an overfished condition; OR adequate information on abundance or biomass is not available High: Abundance or biomass is >125% of BMSY or similar proxy. Points of Adjustment (multiple selections allowed) The population is declining over a generational time scale (as indicated by biomass estimates or standardized CPUE). The abundance in many Sardine populations shows considerable inter-annual variability. In the last decade, an overall decline has been observed in northern Africa (FAO 2007), while recent increases have been observed in southern Africa (FAO 2007). In the Bay of Biscay and Iberian seas abundance has been variable over the last decade (ICES 2010a) Age, size or sex distribution is skewed relative to the natural condition (e.g., truncated size/age structure or anomalous sex distribution) Species is listed as "overfished" OR species is listed as "depleted", "endangered", or "threatened" by recognized national or international bodies Current levels of abundance are likely to jeopardize the availability of food for other species or cause substantial change in the structure of the associated food web. Sardines feed on zooplankton and phytoplankton (Palomera et al. 2007, ICES 2010a). They are important prey for higher tropic levels, including many fish and marine mammals, and play an important role in the transfer of energy up the food web (Palomera et al. 2007, ICES 2010a). Therefore, a low of abundance of Sardines could potentially impact the structure of the food web (Coll et al. 2008, 2009) The population is increasing over a generational time scale (as indicated by biomass estimates or standardized CPUE) Age, size or sex distribution is functionally normal. Age distributions for the Iberian and Biscay Sea region show no evidence of truncation since the early 1980s (ICES 2010b). There is also no evidence of truncation in the length distributions for the African populations (FAO 2007) Species is close to virgin biomass.

6 +0.25 Current levels of abundance provide adequate food for other predators or are not known to affect the structure of the associated food web Points for Abundance HABITAT QUALITY AND FISHING GEAR IMPACTS Core Points (only one selection allowed) Select the option that most accurately describes the effect of the fishing method upon the habitat that it affects 1.00 The fishing method causes great damage to physical and biogenic habitats (e.g., cyanide; blasting; bottom trawling; dredging) The fishing method does moderate damage to physical and biogenic habitats (e.g., bottom gillnets; traps and pots; bottom longlines) The fishing method does little damage to physical or biogenic habitats (e.g., hand picking; hand raking; hook and line; pelagic long lines; mid-water trawl or gillnet; purse seines). Total annual catch for Sardines in the eastern Atlantic has averaged around 1,000,000 tons for the last decade (FAO 2010). In 2010, approximately 100,000 tons of Sardine were landed in the Bay of Biscay and Iberian Seas. Catch off the African coast was around 700,000 tons in 2006 (the last reported year). Sardines are primarily caught with purse seines and pelagic trawls (FAO 2010, Palomera et al. 2007, ICES 2010a). Other minor gears include gillnets, beach seines and trap nets (FAO 2010). Since pelagic gears that have no contact with sea floor are most often used, a score of 3 was awarded. Points of Adjustment (multiple selections allowed) Habitat for this species is so compromised from non-fishery impacts that the ability of the habitat to support this species is substantially reduced (e.g., dams; pollution; coastal development) Critical habitat areas (e.g., spawning areas) for this species are not protected by management using time/area closures, marine reserves, etc.

7 -0.25 No efforts are being made to minimize damage from existing gear types OR new or modified gear is increasing habitat damage (e.g., fitting trawls with roller rigs or rockhopping gear; more robust gear for deep-sea fisheries) If gear impacts are substantial, resilience of affected habitats is very slow (e.g., deep water corals; rocky bottoms) Habitat for this species remains robust and viable and is capable of supporting this species. Sardines are found in the coastal, pelagic eastern Atlantic from 25 to 55 meters. Spawning occurs along the coast in cold waters of C and juveniles are often found in bays (Palomera et al. 2007). There is no indication that there is not enough viable habitat to support this species, therefore points are added Critical habitat areas (e.g., spawning areas) for this species are protected by management using time/area closures, marine reserves, etc. In the eastern Atlantic, fishing is prohibited in the bays and estuaries of Spain and temporary area closures (30-90 days) are used in both Spain and Portugal during the winter months (ICES 2010b). However, since closures are only used for some areas, points are not added Gear innovations are being implemented over a majority of the fishing area to minimize damage from gear types OR no innovations necessary because gear effects are minimal. Since pelagic gears are used in the Sardine fishery (Palomera et al. 2007, ICES 2010a), gear innovations are not necessary to prevent habitat damage If gear impacts are substantial, resilience of affected habitats is fast (e.g., mud or sandy bottoms) OR gear effects are minimal Points for Habitat Quality and Fishing Gear Impacts

8 MANAGEMENT Core Points (only one selection allowed) Select the option that most accurately describes the current management of the fisheries of this species Regulations are ineffective (e.g., illegal fishing or overfishing is occurring) OR the fishery is unregulated (i.e., no control rules are in effect) Management measures are in place over a major portion over the species' range but implementation has not met conservation goals OR management measures are in place but have not been in place long enough to determine if they are likely to achieve conservation and sustainability goals. For the north eastern Atlantic population (Bay of Biscay and Iberian Seas), management advice is provided to Spain and Portugal by the International Council for the Exploration of the Sea (ICES). Spain and Portugal manage the Sardine fishery separately through minimum size limits, maximum daily catch, days at sea restrictions, and closed areas. Portugal also set a total allowable catch for Fishing mortality decreased from 1998 to 2006, but has increased in recent years despite advice not to increase fishing (ICES 2010a). Spawning stock biomass had declined in recent years due to poor recruitment (ICES 2010a). In the past, poor recruitment combined with high fishing mortality has led to periods of minimal spawning stock biomass; therefore is it important that fishing mortality is not increased (ICES 2010b). A single international management plan is needed (ICES 2010a). Management advice for Sardine off the African coast is provided by the Fishery Committee for the Eastern Central Atlantic (CECAF). Commercial catch data, biological data, and acoustic surveys are used to analytically assess the fishery (FAO 2007). References points for this population have been established and a total allowable catch is used to limit Sardine catch (FAO 2007). Fishing in the northern area greatly exceeds the recommended limit and the stock is overexploited, whereas in the southern region the fishery is not fully exploited (FAO 2007). In all of these cases it is unclear whether the current management measures in place will be able to meet sustainability goals; therefore a score of 2 was awarded Substantial management measures are in place over a large portion of the species range and have demonstrated success in achieving conservation and sustainability goals. Points of Adjustment (multiple selections allowed) There is inadequate scientific monitoring of stock status, catch or fishing effort.

9 -0.25 Management does not explicitly address fishery effects on habitat, food webs, and ecosystems. ICES, the management unit for the northeast Atlantic, addresses how the Sardine fishery may impact the ecosystem, in reference to bycatch of non-target species, and there are also ongoing investigations into the role that Sardines plays in the food web (ICES 2010a). However, the management units for African Sardine fisheries do not address these types of ecosystem effects This species is overfished and no recovery plan or an ineffective recovery plan is in place Management has failed to reduce excess capacity in this fishery or implements subsidies that result in excess capacity in this fishery There is adequate scientific monitoring, analysis and interpretation of stock status, catch and fishing effort. Throughout the eastern Atlantic acoustic biomass surveys along with commercial catch data are used in analytical assessments to determine the state of Sardine populations (ICES 2010a, FAO 2007, Bigot and Roos 2009, Bellido et al. 2009, Santojanni and Cingolani 2009, Quintanilla et al. 2009, Basilone and Mazzola 2009, Giannuolaki et al. 2009). Biological reference points have been established for the Sardine populations off the African coasts (FAO 2007), but still need to be established for the other Sardine populations. Points are not added since additional information and analyses are still needed Management explicitly and effectively addresses fishery effects on habitat, food webs, and ecosystems This species is overfished and there is a recovery plan (including benchmarks, timetables and methods to evaluate success) in place that is showing signs of success OR recovery plan is not needed Management has taken action to control excess capacity or reduce subsidies that result in excess capacity OR no measures are necessary because fishery is not overcapitalized Points for Management

10 BYCATCH Core Points (only one selection allowed) Select the option that most accurately describes the current level of bycatch and the consequences that result from fishing this species. The term, "bycatch" used in this document excludes incidental catch of a species for which an adequate management framework exists. The terms, "endangered, threatened, or protected," used in this document refer to species status that is determined by national legislation such as the U.S. Endangered Species Act, the U.S. Marine Mammal Protection Act (or another nation's equivalent), the IUCN Red List, or a credible scientific body such as the American Fisheries Society Bycatch in this fishery is high (>100% of targeted landings), OR regularly includes a "threatened, endangered or protected species." 2.00 Bycatch in this fishery is moderate (10-99% of targeted landings) AND does not regularly include "threatened, endangered or protected species" OR level of bycatch is unknown. Many pelagic fisheries throughout the eastern Atlantic target multiple species. The Spain fisheries target anchovy, mackerel, and horse-mackerel, along with Sardine (ICES 2010a). Portugal fisheries primarily targets Sardine, but chub mackerel, horse-mackerel, and anchovy are also caught (ICES 2010a). In the eastern North Atlantic, the purse seine Sardine fishery is considered to be highly selective and low levels of bycatch have been reported (ICES 2010a). The primary concern for purse seine fisheries is the incidental catch of dolphins (FAO 2010). In the northeastern Atlantic, the purse seine fishery is thought to have a low impact on the common dolphin population (ICES 2010a). The amount of non-targeted and undersized fish that are caught in the African sardine fisheries is unknown Bycatch in this fishery is low (<10% of targeted landings) and does not regularly include "threatened, endangered or protected species." Points of Adjustment (multiple selections allowed) Bycatch in this fishery is a contributing factor to the decline of "threatened, endangered, or protected species" and no effective measures are being taken to reduce it Bycatch of targeted or non-targeted species (e.g., undersize individuals) in this fishery is high and no measures are being taken to reduce it. For the eastern Atlantic, discard estimates have not been quantified, but may be highly sporadic ranging from 0 to 100% (ICES 2010b). Some slipping is known to occur, in which the entire catch is released in the water, rather than being brought on board; this

11 typically occurs when high amounts of undersized or unmarketable bycatch are captured (ICES 2010b). Since, adequate information on bycatch is lacking points are not subtracted Bycatch of this species (e.g., undersize individuals) in other fisheries is high OR bycatch of this species in other fisheries inhibits its recovery, and no measures are being taken to reduce it. The amount of Sardine caught in other fisheries is unknown, but they are likely caught in other pelagic fisheries and may also be caught in bottom trawl fisheries. Due to a lack of information points are not subtracted The continued removal of the bycatch species contributes to its decline Measures taken over a major portion of the species range have been shown to reduce bycatch of "threatened, endangered, or protected species" or bycatch rates are no longer deemed to affect the abundance of the "protected" bycatch species OR no measures needed because fishery is highly selective (e.g., harpoon; spear) There is bycatch of targeted (e.g., undersize individuals) or non-targeted species in this fishery and measures (e.g., gear modifications) have been implemented that have been shown to reduce bycatch over a large portion of the species range OR no measures are needed because fishery is highly selective (e.g., harpoon; spear) Bycatch of this species in other fisheries is low OR bycatch of this species in other fisheries inhibits its recovery, but effective measures are being taken to reduce it over a large portion of the range The continued removal of the bycatch species in the targeted fishery has had or will likely have little or no impact on populations of the bycatch species OR there are no significant bycatch concerns because the fishery is highly selective (e.g., harpoon; spear) Points for Bycatch

12 REFERENCES Alheit J and Hagen E (1997) Long term climate forcing of European herring and sardine populations. Fisheries Oceanography 6: Amenzoui K,, Ferhan-Tachinante F, Yahyaoui A, Kifani S, and Mesfioui AH. Analysis of the cycle of reproduction of Sardina pilchardus off the Moroccan Atlantic coast. Comptes Rendus Biologies 329: Basilone G and Mazzola S (2009) Sardine (Sardina pilchardus) in GSA 16 (South of Sicily). General Fisheries Commission for the Mediterranean (GFCM) Scientific Advisory Committee Twelfth Session. Budva, Montenegro January Bellido JM, Quintanilla L, Torres P, Giraldez A, Ceruso C, Alemany F, and Iglesias M (2009) Sardine (Sardina pilchardus) in GSA 06 (Northern Spain). General Fisheries Commission for the Mediterranean (GFCM) Scientific Advisory Committee 11th Session. Malaga, Spain, Nov. 3- Dec Bigot JL and Roos D. (2009) Sardine (Sardina pilchardus) in GSA 07 (Gulf of Lions). General Fisheries Commission for the Mediterranean (GFCM) Scientific Advisory Committee Twelfth Session. Budva, Montenegro January Catalan IA, Olivar MP, Palomera I, and Berdalet E. (2006) Link between environmental anomalies, growth and condition of pilchard Sardina plichardus larvae in the northwestern Mediterranean. Marine Ecology Progress Series 307: Coll M, Palomera I, Tudela S, and Dowd M (2008) Food-web dynamics in the South Catalan Sea ecosystem (NW Mediterranean) for Ecological Modelling 217: Coll M, Santojanni A, Palomera I, and Arneri E (2009) Food-web changes in the Adriatic Sea over the last three decades. Marine Ecology Progress Series 381: FAO (2010) Species Fact Sheets: Sardina pilchardus. Fisheries and Aquaculture Department, FAO. FAO (2007). FAO working group on the assessment of small pelagic off northwest Africa. FAO Fisheries Report No Agadir, Morocco, April Fishbase (2010) Sardina pilchardus. Ganias K (2009) Linking sardine spawning dynamic to environmental variability. Estuarine Coastal and Shelf Science 84: Ganias K (2008) Ephemeral spawning aggregations in the Mediterranean Sardine, Sardina pilchardus: a comparison with other multiple spawning clupeiods. Marine Biology 155:

13 General Fisheries Commission for the Mediterranean (GFCM) (2006) Marine Resource Fact Sheet: Sardine-Western Part of South Adriatic Sea. General Fisheries Commission for the Mediterranean (GFCM) Scientific Advisory Committee (2009) Report of the transversal workshop on selectivity improvements and bycatch reductions. Tunis, Tunisia, September Giannoulaki M, Somarakis S, Machias A, Kallianiotis A, Siapatis A, and Papaconstantinou C (2009) Sardine (Sardina pilchardus) in GSA 22 (Aegean Sea-NW part). General Fisheries Commission for the Mediterranean (GFCM) Scientific Advisory Committee Twelfth Session. Budva, Montenegro January Guisande C, Cabanas JM, Vergara AR, Riveiro I (2001) Effect of climate on recruitment success of Atlantic Iberian Sardine Sardina pilchardus. Marine Ecology Progress Series 223: Guisande C, Vergara AR, Riveiro I, Cabanas JM (2004) Climate change and abundance of the Atlantic-Iberian Sardine. Fisheries Oceanography 13: ICES (2010a) Sardine in Divisions VIIIc and IXa (Bay of Biscay and Western Iberian Seas). ICES Advice ICES (2010b) Report of the Working Group on Anchovy and Sardine (WGANSA), June 2010, Vigo, Spain. ICES CM 2010/ACOM: pp. Morello EB and Arneri E (2009) Anchovy and Sardine in the Adriatic Sea- An ecological review. Oceanography and Marine Biology 47: Omar KADA and My Hachem IDRISSI (2009) Sardine (Sardina pilchardus) in GSA 03 (Southern Alboran Sea). General Fisheries Commission for the Mediterranean (GFCM) Scientific Advisory Committee 11th Session. Malaga, Spain, Nov. 3- Dec Palomera I, Olivar MP, Salat J, Sabates A, Coll M, Garcia A, and Morales-Nin B (2007) Small pelagic fish in the NW Mediterranean Sea: An ecological review. Progress is Oceanography 74: Parrish RH, Serra. R, and Grant WS (1989) The monotypic sardines, Sardina and Sardinops: their taxonomy, stock structure, and zoogeography. Can. J. Fish. Aquat. Sci. 46~ Quintanilla LF, Bellido JM, Torres P, Giraldez A, Ceruso A, Alemany F, and Iglesias M (2009) Sardine (Sardina pilchardus) in GSA 01 (Northern Alboran Sea). General Fisheries Commission for the Mediterranean (GFCM) Scientific Advisory Committee 11th Session. Malaga, Spain, Nov. 3- Dec Santojanni A and Cingolani N (2009) Sardine (Sardina pilchardus) in GSA 17 (Northern Adriatic). General Fisheries Commission for the Mediterranean (GFCM) Scientific Advisory Committee 11th Session. Malaga, Spain, Nov. 3- Dec

14 Silva A, Carrera P, Masse J, Uriate A, Santos MB, Oliveira PB, Soares E, Porterio C, and Stratoudakis Y (2008) Geographic variability of sardine growth across the northeastern Atlantic and Mediterranean Sea. Fisheries Research 90: Silva A, Santos MB, Caneco B, Pestana G, Porteiro C, Carreca P, Stratoudakis Y (2006) Temporal and geographic variability of sardine maturity at length in the northeastern Atlantic and the western Mediterranean. ICES Journal of Marine Science 63: Solari AP, Santamaria MTG, Borges MF, Santos AMP, Mendes H, Balguerias E, Castro JJ, Bas C. (2010) On the dynamics of Sardine Pilchardus: orbits of stability and environmental forcing. ICES Journal of Marine Science 67: Somarakis S, Ganias K, Siapatis A, Koutsikopoulos C, Machias A, Papaconstantinou C (2006) Spawning habitat and daily egg production of sardine (Sardina pilchardus) in the eastern Mediterranean. Fisheries Oceanography, 15 (4): Stratoudakis Y, Coombs S, Lago de Lanzós A, Halliday N, Costas G, Caneco B, Franco C, Conway D, Santos MB, Silva A, Bernal M (2007) Sardine (Sardina pilchardus) spawning seasonality in European waters of the northeast Atlantic. Marine Biology 152: Tsagarakis K, Machias A, Somarakis S, Giannoulaski M, Palialexis A, Valavanis VD (2008) Habitat discrimination of juvenile sardines in the Aegean Sea using remotely sensed environmental data. Hydrobiologia 612:

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