The European pilchard (Sardina pilchardus) is a species of ray-finned fish in the monotypicgenusSardina. The young of the species are among the many fish that are sometimes called sardines.[3][4] This common species is found in the northeast Atlantic, the Mediterranean, and the Black Sea at depths of 10–100m (33–328ft).[1] It reaches up to 27.5cm (10.8in) in length and mostly feeds on planktoniccrustaceans.[2] This schooling species is a batch spawner where each female lays 50,000–60,000 eggs.[2]
The European pilchard is a small to medium-sized, somewhat elongated, herring-like fish. The origin of the pelvic fins is well behind that of the dorsal fin, and the last two soft rays on the anal fin are larger than the remainder. The upper parts are green or olive, the flanks are golden and the belly is silvery.[3]
Not to be confused with its American counterpart, the California sardine, Sardina sagax, the European sardine S. pilchardus does not have a row of dark blotches. They also have scales that are in an irregular order. Ventral scutes on the European species have sturdier keels that are also more keeled. Gillrakers are significantly different from those on Sardina sagax.[5]
Distribution and habitat
The European pilchard occurs in the northeastern Atlantic Ocean, the Mediterranean Sea and the Black Sea. Its range extends from Iceland and the southern part of Norway and Sweden southwards to Senegal in West Africa. In the Mediterranean Sea it is common in the western half and the Adriatic Sea, but uncommon in the eastern half and the Black Sea.[6] It is a migratory, schooling, largely coastal species but sometimes travels as far as 100km (62mi) out to sea. During the day it is mostly in the depth range 25 to 55m (80 to 180ft) but can go as deep as 100m (330ft). At night it is generally from 10 to 35m (33 to 115ft) beneath the surface.[1]
Taxonomy
S. pilchardus is part of the Clupeidae family along with over 300 other species. Also known as the North Atlantic-Mediterranean sardine, it was the first sardine to be described. It was originally put under Clupea from 1842 to 1879. Under this genus, Gunther (1868) gave a general description of a terminal mouth, minute or absent teeth, a complete midventral series of scutes, and an anal fin. However, Regan (1916,1917) divided the genus into two subgenus: the subtropical Sardina and tropical Sardinella. S. pilchardus and two other species (e.g. S. sagax and S. neopilchardus) were put under Sardina. Between the two, gill raker counts and head length distinctions differentiate each genus.
Svetovidov (1952) noted common characteristics within the Clupidae family which include: a more anterior dorsal fin, a more developed lateral line canal in the head, last two fin rays of the anal fin are elongated lobes, dark lateral pigment spots, elongated scales on the base of caudal fin lobes, interrupted transverse striations on body scales, and radial striations on the operculum of Sardina and Sardinops. A distinguishing feature of the Sardina from other genera is the body scales, which vary in size. Smaller scales are hidden underneath the larger scales.[5] Otolith variations are most likely attributed to environmental differences as opposed to genetic variation in S. pilchardus.[7]
Ecology
Juveniles schooling by day in shallow water in the Ligurian Sea, Italy
In the Mediterranean, the European pilchard moves offshore in the autumn, preferring the deeper cooler waters and constant salinity out at sea to the variable temperatures and salinities of inshore waters. Spawning for S. pilchardus occurs in open waters with long larval periods. S. pilchardus is a capital breeder — it stores energy during a specific period (i.e. spring-summer) and then uses it for reproduction.[7] Spawning starts to take place in winter, and in early spring, juveniles, larvae and some adults move towards the coast, while other adults migrate inshore later in the year. Multiple batches of eggs are produced over a long breeding period, total fecundity being 50,000 to 60,000. Most juveniles become sexually mature at about a year old and a length of 13 to 14cm (5.1 to 5.5in); pilchards are fully grown at about 21cm (8.3in) when aged about eight years.[8]
The diet consists of both zooplankton and phytoplankton. The zooplankton is largely copepods and their larvae, which make daily vertical migrations to feed near the surface at night, and this is when the adult pilchards feed on them; juveniles feed during the day as well.[8] Along with the European anchovy (Engraulis encrasicolus), the European pilchard plays an important intermediate role in the Mediterranean ecosystem as a consumer of plankton and as a food for larger demersal predators such as the European hake (Merluccius merluccius) and the European conger eel (Conger conger). This role is particularly noticeable in the Adriatic Sea where the water is shallow, the food chain is shorter and energy is retained within the basin; overfishing of pilchard and anchovy can thus cause dramatic changes in the ecosystem.[8]
S. pilchardus is a short-lived, fast-maturing, and highly fecund species. At both the larval and adult stages of its life history, it has a high ability to disperse which results in high gene flow and thus low genetic differentiation between subareas of the Mediterranean.[7]
Nematodes found in S. pilchardus include the Anisakidae and Raphidascarididae. However, despite being a widely eaten fish around Europe, especially in the Mediterranean, only eight cases of human anisakidosis have been reported since 1991. A study by Fuentes et al. (2022) show that the consumption of S. pilchardus poses an unlikely health hazard for parasite infection of Anisakis spp., even when consumed raw or lack of adequate preparation through freezing. But as with all fish consumed raw, proper precautions should be taken, such as consumption of sardines caught in the Mediterranean and smaller specimens. Additionally, consultation of labels and traceability data can help to further minimize risk.[9]
Fisheries and uses
European pilchards, center right, are the cheapest thing for sale at this fish stall in Syracuse, SicilyGlobal capture production of European pilchard (Sardina pilchardus) in million tonnes from 1950 to 2022, as reported by the FAO
There are important fisheries for this species in most of its range. It is mainly caught with purse seines and lampara nets, but other methods are also used including bottom trawling with high opening nets. S. pilchardus is one of the most heavily impacted species impacted by fisheries in the Mediterranean region.[11] In a 2022 report from the Fishery Committee for The Eastern Central Atlantic (CECAF), S. pilchardus attributed to 48% of overall small pelagic species catch, or the equivalent of 2.3 million tons,[12] with Morocco, Portugal and Spain having the largest catches. As such, S. pilchardus has a huge economic impact on the surrounding land regions. The Food and Agriculture Organization (FAO) considers the Moroccan fishery overfished.[6]
Globally, Sardina pilchardus ranks number eight for most captured species. However, recently, sardines in the Mediterranean regions have shown lower growth, fecundity, and spawning rates than those in Atlantic regions. This has been possibly attributed to the oligotrophy of the Mediterranean. In addition, a decreasing trend of S. pilchardus biomass has occurred. A possible explanation could be that the disappearance of older fish due to a less-than-ideal environment and fishing pressure could be the cause of declining biomass of S. pilchardus.[11]
Little is known about status of the stock of S. pilchardus despite the economic dependency on the surrounding nations, particularly in the West African region [9]. Some research has been conducted to determine if there are distinct genetic and morphological differences between certain stocks within their range. According to Baldé et al. (2022), "plasticity in the growth performance, survival, and other life-history characteristics […] is the key to their dynamics." They are sensitive to environmental changes and are good indicators of how climate change is impacting marine environments.[13]
The adults may be sold as pilchards; the juveniles, as sardines.[3] The terms "sardine" and "pilchard" are not precise, and what is meant depends on the region. The United Kingdom's Sea Fish Industry Authority, for example, classifies sardines as young pilchards.[14] One criterion suggests fish shorter in length than 15cm (6in) are sardines, and larger fish are pilchards.[15] The FAO/WHO Codex standard for canned sardines cites 21 species that may be classed as sardines.[4]
Xouba is a small version of the pilchard which is prevalent in Spain. Xoubas, are small sardines, and a they come from Galicia, Spain.[16][17] In comparison to the other sardines, the xouba, is diminutive.[18] Xouba means little fish; the fish lives is shallow areas of water between 5m (16ft) and 50m (160ft).[18]
The fish is sold fresh, frozen or canned, or is salted and smoked or dried; as the flesh is of low value, some of the catch is used for fishing bait or fertilizer and some is manufactured into fish meal.[1]
Genetics
Mitochondrial sequence data have shown evidence of an early genetic bottleneck in Safi and a significant genetic differentiation within the sardine population.[19] However, this has been challenging for researchers to definitively claim due to large population sizes and the sardine’s high dispersal ability. Additionally, the lack of oceanographic barriers allow for further gene flow between populations.
Eight micro-satellite loci showed polymorphism and no significant linkage disequilibrium.[20] Due to possible homozygote excess and null alleles, most loci showed differences from the Hardy-Weinberg Equilibrium. Further tests showed that the S. pilchardus populations from across nine locations over the Atlantic Ocean and Mediterranean Sea act as one evolutionary unit. Slight genetic differences found could be attributed to isolation by distance. Thus, there is support for two distinct groups via mitochondrial data, while micro-satellite data supports weak genetic structure within a evolutionary unit.[19]
Mitochondrial and micro-satellite analyses of the Sp2, Sp7, Sp8 and Sp15 loci with biologic parameter and genetic analysis show homogeneity in the Atlantic Moroccan coast population of S. pilchardus. In Baibai et al. (2017)’s biologic and morphometric analysis, they found high variability within the stocks of the Moroccan Atlantic, with two distinct morphological groups that are geographically separated in the Northwest African Coast and South region of Morocco. Specifically, northern Moroccan sardines have a smaller head to body ratio than those in the south. These are in addition to the established differences in the position of the pelvic and anal fin between the same groups. The morphological differences, however, are not genetically based. Instead, it is the result of phenotypic plasticity, a trait known to help sardine stocks survive in the different waters that they inhabit.[20]
A genetic analysis of relevant microsatellite loci and mitochondrial analysis further support the claim of non-genetically based phenotypic difference. All four loci studied were analyzed to be independent of each other. Of note, loci Sp2 and Sp8 carried a high number of private alleles, although this is consistent with other marine fishes. These same two loci are responsible for the small genetic structure found in S. pilchardus. Due to homozygote excess and null alleles, significant values of the inbreeding coefficient, Fis, was found in Safi, Malaga, and Cadiz. Mitochondrial analysis showed significant genetic structure compared to previous analyses with significant nucleotide differences and higher nucleotide diversity in Cap Blanc and Galician samples. Possible explanations for the isolation of certain populations could be explained by gyres or upwelling, such as that in south Morocco, along with distance.[20]
Stocks
In a study by Huertas et al. (2023), S. pilchardus somatic condition, relative condition, hepasomatic indices, and reproductive condition were used to compare four sub areas of the Mediterranean: Northern Alboran, Northern Spain, Northern Adriatic, and Aegean Sea. The study found that there are differences in energy storage that are supported by underlying genetic differences. Thus, there is heterogeneity in regard to reproduction.
Northern Spain showed the lowest somatic condition patterns. This subarea has also had a decline in older fish which limits the plasticity of sardines to environmental changes and thus affects spawn quality. However, the Northern Adriatic stock has body mass overall above that of the other stocks. This could be due to year-round accessibility to feeding resources year-round. Furthermore, the main phytoplankton bloom in this area coincides with sardine reproductive season. But phytoplankton still only accounts for around 10% of its diet, with copepods being the majority (56%). The study found that because reproduction is dependent availability of resources, the Northern Adriatic population’s storage period is not as distinctive as other stocks.[21]
Northern Alboran stock was recorded to show earlier gonad maturation in the summer than their Atlantic neighbors. The authors speculate individuals from the Atlantic swim into the Mediterranean, spawn, and the drop in temperature along the Alboran coast favors sardine nursery grounds. The Northern Alboran stock had slightly more energy stored in the muscles compared to the other groups studied. Additionally, only the Northern Alboran area saw improvement in body condition from 2010 while other subareas saw a dramatic decline in body condition and maximum size between 1975–2015. The authors speculate there is possible overlap between the Atlantic and Alboran individuals which causes genetic variability and plasticity. Moreover, this area is more favorable to those with an affinity for cool temperatures with less seasonal variation. The authors urge for close monitoring of the Northern Spain sardine population due to rapid environmental change vulnerability and heavy fishing pressures.[21]
In a more recent study, analysis of the genome of the European sardine S. pilchardus and mitochondrial data revealed evidence of at least three genetic clusters within the natural range (Eastern Mediterranean to the Azores archipelago). However, more research will be needed to more firmly establish these findings.[22]
Atlantic herring is a herring in the family Clupeidae. It is one of the most abundant fish species in the world. Atlantic herrings can be found on both sides of the Atlantic Ocean, congregating in large schools. They can grow up to 45 centimetres (18 in) in length and weigh up to 1.1 kilograms (2.4 lb). They feed on copepods, krill and small fish, while their natural predators are seals, whales, cod and other larger fish.
Sardine and pilchard are common names for various species of small, oily forage fish in the herring suborder Clupeoidei. The term 'sardine' was first used in English during the early 15th century; a somewhat dubious etymology says it comes from the Italian island of Sardinia, around which sardines were once supposedly abundant.
The albacore, known also as the longfin tuna, is a species of tuna of the order Scombriformes. It is found in temperate and tropical waters across the globe in the epipelagic and mesopelagic zones. There are six distinct stocks known globally in the Atlantic, Pacific, and Indian oceans, as well as the Mediterranean Sea. The albacore has an elongate, fusiform body with a conical snout, large eyes, and remarkably long pectoral fins. Its body is a deep blue dorsally and shades of silvery white ventrally. Individuals can reach up to 1.4 m in length.
The Atlantic bluefin tuna is a species of tuna in the family Scombridae. It is variously known as the northern bluefin tuna, giant bluefin tuna, and formerly as the tunny.
The Atlantic mackerel, also known as Boston mackerel, Norwegian mackerel, Scottish mackerel or just mackerel, is a species of mackerel found in the temperate waters of the Mediterranean Sea, the Black Sea, and the northern Atlantic Ocean, where it is extremely common and occurs in huge shoals in the epipelagic zone down to about 200 m (660 ft). It spends the warmer months close to shore and near the ocean surface, appearing along the coast in spring and departing with the arrival of colder weather in the fall and winter months. During the fall and winter, it migrates out into deeper and more southern water, seeking warmer temperatures.
The ciscoes are salmonid fish that differ from other members of the genus Coregonus in having upper and lower jaws of approximately equal length and high gill raker counts. These species have been the focus of much study recently, as researchers have sought to determine the relationships among species that appear to have evolved very recently. Cisco is also specifically used for the North American species Coregonus artedi, also known as lake herring.
The Alboran Sea is the westernmost portion of the Mediterranean Sea, lying between the Iberian Peninsula and the north of Africa. The Strait of Gibraltar, which lies at the west end of the Alboran Sea, connects the Mediterranean with the Atlantic Ocean.
The Dolly Varden trout is a species of salmonid ray-finned fish native to cold-water tributaries of the Pacific Ocean in Asia and North America. Despite the name "trout", it belongs to the genus Salvelinus (chars), which includes 51 recognized species, the most prominent being the brook, lake and bull trout as well as the Arctic char. Although many populations are semi-anadromous, riverine and lacustrine populations occur throughout its range. It is considered by taxonomists as part of the Salvelinus alpinus complex, as many populations of bull trout, Dolly Varden trout and Arctic char overlap.
The European sprat, also known as brisling, brisling sardine, bristling, garvie, garvock, Russian sardine, russlet, skipper or whitebait, is a species of small marine fish in the herring family Clupeidae. Found in European, West Asian and North African waters, it has silver grey scales and white-grey flesh. Specific seas in which the species occurs include the Irish Sea, Black Sea, Baltic Sea and Sea of the Hebrides. The fish is the subject of fisheries, particularly in Scandinavia, and is made into fish meal, as well as being used for human consumption. When used for food it can be canned, salted, breaded, fried, boiled, grilled, baked, deep fried, marinated, broiled, and smoked.
The slender rainbow sardine is a small, subtropical, salt water fish of the Indian Ocean and the South China Sea which was first described by Bleeker in 1849. Dussumieria hasselti and Dussumieria productissima are synonyms for this same fish. It is a round herring of the family Clupeidae.
The greater amberjack, also known as the allied kingfish, great amberfish, greater yellowtail, jenny lind, Sea donkey, purplish amberjack, reef donkey, rock salmon, sailors choice, yellowtail, and yellow trevally, is a species of predatory ray-finned fish in the family Carangidae, the jacks and pompanos. It is found in temperate, subtropical, and tropical seas around the world. It is a popular quarry species for recreational fisheries and is important in commercial fisheries. It is the largest species in the family Carangidae.
The European anchovy is a forage fish somewhat related to the herring. It is a type of anchovy; anchovies are placed in the family Engraulidae. It lives off the coasts of Europe and Africa, including in the Mediterranean Sea, the Black Sea, and the Sea of Azov. It is fished by humans throughout much of its range.
An anchovy is a small, common forage fish of the family Engraulidae. Most species are found in marine waters, but several will enter brackish water, and some in South America are restricted to fresh water.
Sardinella zunasi is a species of ray-finned fish in the family Clupeidae, the herrings and sardines. It is native to the northwestern Pacific Ocean, where it occurs near shore along the Asian coastlines from southern Japan to Taiwan.
Merluccius merluccius, the European hake, is a merluccid hake of the genus Merluccius. Other vernacular names include Cornish salmon and herring hake. It is a predatory species, which was often netted alongside one of its favoured prey, the Atlantic herring, hence the latter common name. It is found in the eastern Atlantic from Norway and Iceland south to Mauritania and into the Mediterranean Sea. It is an important species in European fisheries and is heavily exploited, with some populations being fished unsustainably.
The blue ling is a member of the cod family from the North Atlantic. It is usually 70 to 110 cm long, but the maximum length is 155 cm. Blue ling feed on fish and crustaceans and benthic invertebrates. The fish reaches sexual maturity at the age of six to 12 years.
Sardines ("pilchards") are a nutrient-rich, small, oily fish widely consumed by humans and as forage fish by larger fish species, seabirds and marine mammals. Sardines are a source of omega-3 fatty acids. Sardines are often served in cans, but can also be eaten grilled, pickled, or smoked when fresh.
The Alosidae, or the shads, are a family of clupeiform fishes. The family currently comprises four genera worldwide, and about 32 species.
Ichthyodinium is a monotypic genus of dinoflagellates in the family Dinophysaceae. Ichthyodinium chabelardi (/ɪkθioʊˈdɪniəm/) is currently the sole described species of the genus.
Xouba also known as little sardine or parrocha is a small sardine and a type of Pelagic fish. Xoubas are small sardines from Galicia, Spain. They are a species of ray-finned fish in the monotypic genus Sardina. The juvenile Xouba has the scientific name of European pilchard which is in the family Clupeidae.
1 2 Parrish, R.H.; Serra, R.; Grant, W.S. (1989). "The Monotypic Sardines, Sardina and Sardinops: Their Taxonomy, Distribution, Stock Structure, and Zoogeography". Canadian Journal of Fisheries and Aquatic Sciences. 46: 2019–2036.
1 2 3 Jemaa, S.; Bacha, M.; Khalaf, G.; Dessailly, D.; Rabhi, K.; Amara, R. (2015). "What can otolith shape analysis tell us about population structure of the European sardine, Sardina pilchardus, from Atlantic and Mediterranean waters?". Journal of Sea Research. 96: 11–17.
↑ Fuentes, M.; Madrid, E.; Melia, L..V.; Casafe, F.; Saez-Duran, S.; Trelis, M.; Debenedetti, A.L. (2022). "Nematode Parasites of the European Pilchard, Sardina pilchardus (Walbaum, 1792): A Genuine Human Hazard?". Animals. 12 (15): 1877.
1 2 Ferhani, K.; Bensmail, S.; Guechaoui, M.; Bennoui, A.; Chabet Dis, C. (2024). "Age and Growth of European Pilchard Sardina pilchardus (Clupidae) from the Southwestern Mediterranean Sea". Journal of Ichthyology.
↑ Kindong, R.; Sarr, O.; Achille Pandong, N.; Wu, F.; Zhu, J.; Dai, X. (2024). "Towards sustainable exploitation of European pilchard (Sardina pilchardus) stocks off West Africa waters". Environmental and Sustainability Indicators. 23: 100460.
↑ Balde, B.S.; Brehmer, P.; Faye, S.; Diop, P. (2024). "Population Structure, Age, and Growth of Sardine (Sardina pilchardus, Walbaum, 1792) in an Upwelling Environment". Fishes. 7 (178).
↑ "FAQs". Seafish. Archived from the original on 22 July 2011. Retrieved 22 February 2010.
1 2 G. Gonzalez, Elena; Zardoya, Rafael (2007). "Relative role of life-history traits and historical factors in shaping genetic population structure of sardines (Sardina pilchardus)". BMC Evolutionary Biology. 7 (197).
1 2 3 Baibai, T.; Oukhattar, L.; Vasquez Quinteiro, J.; Mesfioui, A.; Rey-Mendez, M..; Soukri, A. (2012). "First global approach: morphological and biological variability in a genetically homogenous population of the European pilchard, Sardina pilchardus (Walbaum, 1792) in the North Atlantic coast". Reviews in Biology and Fisheries. 22: 63–80.
1 2 Huertaa, M.C.; Frigola-Tepe, X.; Vinas, J.; Munoz, M. (2023). "From west to east: Heterogeneity in the life history traits of a small pelagic fish (Sardina pilchardus) throughout the Mediterranean". Frontiers in Marine Science. 10: 1187071.
↑ Fonesca, R.; Campos, P.F.; Rey-Iglesia, A.; Barroso, G.V.; Bergeron, L.A.; Nande, M.; Tuya, F.; Abidli, S.; Perez, M.; Riveiro, I.; etal. (2024). "Population Genomics Reveals the Underlying Structure of the Small Pelagic European Sardine and Suggests Low Connectivity within Macaronesia". Genes (Basel). 15 (2): 170.
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