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Pecten maximus

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Pecten maximus
Pecten maximus Pilgermuschel.jpg
Two beachworn upper (flat) valves of Pecten maximus from Wales
Scientific classification edit
Kingdom: Animalia
Phylum: Mollusca
Class: Bivalvia
Order: Pectinida
Family: Pectinidae
Genus: Pecten
Species:
P. maximus
Binomial name
Pecten maximus
Synonyms[1]
  • Ostrea maxima Linnaeus, 1758
  • Pecten medius Daniel, 1883
  • Pecten vulgaris da Costa, 1778
  • Plicatula similis G. B. Sowerby II, 1842

Pecten maximus, common names the great scallop, king scallop, St James shell or escallop, is a northeast Atlantic species of scallop, an edible saltwater clam, a marine bivalve mollusc in the family Pectinidae. This is the type species of the genus. This species may be conspecific with Pecten jacobaeus, the pilgrim's scallop, which has a much more restricted distribution.[2]

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Species

Species

In biology, a species is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. Other ways of defining species include their karyotype, DNA sequence, morphology, behaviour, or ecological niche. In addition, paleontologists use the concept of the chronospecies since fossil reproduction cannot be examined.

Scallop

Scallop

Scallop is a common name that encompasses various species of marine bivalve mollusks in the taxonomic family Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related families within the superfamily Pectinoidea, which also includes the thorny oysters.

Clam

Clam

Clam is a common name for several kinds of bivalve molluscs. The word is often applied only to those that are edible and live as infauna, spending most of their lives halfway buried in the sand of the seafloor or riverbeds. Clams have two shells of equal size connected by two adductor muscles and have a powerful burrowing foot. They live in both freshwater and marine environments; in salt water they prefer to burrow down into the mud and the turbidity of the water required varies with species and location; the greatest diversity of these is in North America.

Pecten jacobaeus

Pecten jacobaeus

Pecten jacobaeus, the Mediterranean scallop, is a species of scallop, an edible saltwater scallop, a marine bivalve mollusc in the family Pectinidae, the scallops.

Description

The shell of Pecten maximus is quite robust and is characterised by having "ears" of equal size on either side of the apex. The right, or lower, valve is convex and slightly overlaps the flat left, or upper, valve, which is flat. Larger specimens have a nearly circular outline and the largest may measure 21 cm in length. The "ears" are prominent and are a minimum of half the width of the shell with the byssal notch situated in the right anterior ear being slight and not serrated. The sculpture of the valves is distinctive and consists of 12 to 17 wide radiating ribs and numerous concentric lines which clearly show the scallop's growth history, while the "ears" show a few thin ribs which radiate from the beaks. The radiating ribs reach the margins of the valves and this creates a crenulated form. The left valve is normally reddish-brown while the right valve varies from white through cream to shades of pale brown contrasting with pink, red or pale yellow tints; either valve may show zigzag patterns and may also show bands and spots of red, pink or bright yellow.[3]

The colour of the body of Pecten maximus is pink or red with the mantle marbled brown and white. When young they are attached to the substrate by a byssus but mature animals are capable of swimming by the opening and rapid closing of the valves. The adductor muscle which is used to close and open the valves is very large and powerful. The foot is a finger-like organ, which spins the byssal threads, which pass through the byssal notch on the ears. The margin of the mantle has two layers: the inner layer is finely fringed while the outer is lined with long tentacles with two series totalling 30–36 dark blue or green simple eyes or ocelli in two rows at their base.[4]

Distribution

Pecten maximus occurs in the eastern Atlantic along the European coast from northern Norway, south to the Iberian peninsula, it has also been reported off West Africa, off the Macaronesian Islands. In Great Britain and Ireland it is distributed all round the coast but it is uncommon and localised on the eastern North Sea coast.[5] It prefers offshore waters down to 100 metres (330 feet; 55 fathoms) depth.[4]

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Norway

Norway

Norway, officially the Kingdom of Norway, is a Nordic country in Northern Europe, the mainland territory of which comprises the western and northernmost portion of the Scandinavian Peninsula. The remote Arctic island of Jan Mayen and the archipelago of Svalbard also form part of Norway. Bouvet Island, located in the Subantarctic, is a dependency of Norway; it also lays claims to the Antarctic territories of Peter I Island and Queen Maud Land. The capital and largest city in Norway is Oslo.

Macaronesia

Macaronesia

Macaronesia is a collection of four volcanic archipelagos in the North Atlantic, off the coasts of Africa and Europe. Each archipelago is made up of a number of Atlantic oceanic islands, which are formed by seamounts on the ocean floor whose peaks have risen above the ocean's surface. Some of the Macaronesian islands belong to Portugal, some belong to Spain, and the rest belong to Cape Verde. Politically, the islands belonging to Portugal and Spain are part of the European Union. Geologically, Macaronesia is part of the African tectonic plate. Some of its islands – the Azores – are situated along the edge of that plate at the point where it abuts the Eurasian and North American plates.

North Sea

North Sea

The North Sea lies between Great Britain, Denmark, Norway, Germany, the Netherlands, Belgium and France. An epeiric sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the Norwegian Sea in the north. It is more than 970 kilometres (600 mi) long and 580 kilometres (360 mi) wide, covering 570,000 square kilometres (220,000 sq mi).

Biology

Live individual of Pecten maximus on the right, next to Ostrea edulis
Live individual of Pecten maximus on the right, next to Ostrea edulis

Pecten maximus frequently creates a slight hollow in the substrate for its shell to lie in by opening and closing the valve to eject water from the mantle cavity, which raises the shell at an angle to the substrate so that subsequent water jets into the sediment and create a recess.[5] Once settled, sand, mud, gravel or living organisms coat the upper valve, and the margin of the shell, with the tentacles and eyes, is all that is visible.[3] They are filter feeders which extract particles from the surrounding water via a feeding current which is drawn by cilia across the gills where the food particles are trapped, then taken to the mouth in a stream of mucous.[6]

Pecten maximus swims but this is generally limited to escape reactions. The main predators which cause this reaction when detected are the mollusc eating starfish Asterias rubens and Astropecten irregularis, although starfish which do not feed on molluscs can cause limited jumping or valve-closing reactions. The swimming action is performed by rapidly clapping the valves and expelling jets of water from each side of the hinge so that it moves with the curved edge of the shell at the front. The scallop jumps forwards by gradually relaxing the adductor muscle and then rapidly opening and closing the valves.[5]

Pecten maximus tends to be more numerous in areas where they are not fully exposed to strong currents. Scallops which live in sheltered habitats grow faster than scallops in areas exposed to wave action, possibly due to the filter feeding apparatus being unable to function because of high concentrations of particulate matter in the water in areas subject to high levels of wave exposure. Another factor that may be significant is that the processes of larval settlement and byssal attachment are rather delicate and would be disturbed in strong currents. Abundance and growth rates are negatively correlated with the amount of mud in the substrate.[3]

Scallops use larval motility to distribute themselves, as the adult scallops have limited mobility. The distribution of the larvae is affected by factors such as local hydrographic regimes and their survival, and this results in the scallops having an aggregated distribution within their geographic range. This means that the major fishing grounds are normally widely separated and each fishing ground's environmental conditions mean there are marked differences in structures of the populations, although the genetics of scallops are rather uniform across its range.[5]

The reproductive cycle of Pecten maximus is extremely variable and the spawning may be influenced by both internal and external factors such as age and temperature respectively but is also influenced by genetic adaptation. Generally, mature scallops spawn over the summer months starting in April or May and lasting to September. They are hermaphroditic and have distinct tongue-shaped gonads which are red or orange in colour for the female gonad and white for the male.[6] It is estimated that a three-year-old scallop releases 15–21 million oocytes per emission. There appear to be two spawnings in many parts of the range, normally there is a partial one in the Spring and a full one in late August, however younger scallops have a single spawning event in the late summer. In some areas this pattern is reversed and the major spawning is in the Spring. After spawning the animals undergo period of recovery of the gonad before they spawn again. Fertilization of the gametes is external and either sperm or oocytes can be released into the water column first.[5]

Since the larval stage of Pecten maximus is relatively long, up to a month, the potential for dispersal is quite high, even smaller adults can use the byssus to drift too. However, in at least some populations, genetic studies show that there is little contribution from more distant populations and that these populations probably sustain themselves.[5]

In waters around the United Kingdom Pecten maximus become sexually mature at around 2–3 years old and when they reach 80 to 90 mm in shell length. Where they are not exploited, they may live for more than 20 years and reach shell lengths of more than 200mm.[7] Scallops in shallow water grow faster than those in deeper water; the growth halts in winter and starts again in spring, producing concentric growth rings which are used to age the scallops.[5]

Genomics

A draft genome is presented by Kenny et al 2020.[8][9] Their assembly is 918Mb, and they estimate a genome size of 1,150Mb, 1.7% heterozygosity, with 27.0% being repeats, in total coding 67,741 genes.[9] Recent (as of 2021) improvements in read length helped Kenny to resolve questions of copy number variation in P. maximus which were previously indecipherable.[8]

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Ostrea edulis

Ostrea edulis

Ostrea edulis, commonly known as the European flat oyster, is a species of oyster native to Europe. In Britain and Ireland, regional names include Colchester native oyster, mud oyster, or edible oyster. In France, Ostrea edulis are known as huîtres plates except for those that come from the Belon River estuary in Brittany, France, which are known as Belons.

Starfish

Starfish

Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea. Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish are also known as asteroids due to being in the class Asteroidea. About 1,900 species of starfish live on the seabed in all the world's oceans, from warm, tropical zones to frigid, polar regions. They are found from the intertidal zone down to abyssal depths, at 6,000 m (20,000 ft) below the surface.

Astropecten irregularis

Astropecten irregularis

Astropecten irregularis is a sea star of the family Astropectinidae. Common names include Sand sea star.

Sperm

Sperm

Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction. Animals produce motile sperm with a tail known as a flagellum, which are known as spermatozoa, while some red algae and fungi produce non-motile sperm cells, known as spermatia. Flowering plants contain non-motile sperm inside pollen, while some more basal plants like ferns and some gymnosperms have motile sperm.

Oocyte

Oocyte

An oocyte, oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female germ cells produce a primordial germ cell (PGC), which then undergoes mitosis, forming oogonia. During oogenesis, the oogonia become primary oocytes. An oocyte is a form of genetic material that can be collected for cryoconservation.

Byssus

Byssus

A byssus is a bundle of filaments secreted by many species of bivalve mollusc that function to attach the mollusc to a solid surface. Species from several families of clams have a byssus, including pen shells (Pinnidae), true mussels (Mytilidae), and Dreissenidae.

Genome

Genome

In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA. The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as regulatory sequences, and often a substantial fraction of 'junk' DNA with no evident function. Almost all eukaryotes have mitochondria and a small mitochondrial genome. Algae and plants also contain chloroplasts with a chloroplast genome.

Genome size

Genome size

Genome size is the total amount of DNA contained within one copy of a single complete genome. It is typically measured in terms of mass in picograms or less frequently in daltons, or as the total number of nucleotide base pairs, usually in megabases. One picogram is equal to 978 megabases. In diploid organisms, genome size is often used interchangeably with the term C-value.

Copy number variation

Copy number variation

Copy number variation (CNV) is a phenomenon in which sections of the genome are repeated and the number of repeats in the genome varies between individuals. Copy number variation is a type of structural variation: specifically, it is a type of duplication or deletion event that affects a considerable number of base pairs. Approximately two-thirds of the entire human genome may be composed of repeats and 4.8–9.5% of the human genome can be classified as copy number variations. In mammals, copy number variations play an important role in generating necessary variation in the population as well as disease phenotype.

Predators and diseases

As well as Asterias rubens and Astropecten irregularis major predators on Pecten maximus are crabs such as Cancer pagurus, Carcinus maenas, Liocarcinus puber and Liocarcinus depurator will prey on the scallops as they grow.[10] The anemone Anthopleura ballii was found preying on young specimens of P.maximus in south-western Ireland.[11]

The larvae of Pecten maximus are attacked by the bacterium Vibrio pectenicida, which was described in 1998 as a new species after incidents of mortality among cultured scallops in France in the early 1990s.[12] Other strains of pathogenic bacteria were detected in Norway following mass mortality of larvae in culture.[13]

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Cancer pagurus

Cancer pagurus

Cancer pagurus, commonly known as the edible crab or brown crab, is a species of crab found in the North Sea, North Atlantic Ocean, and perhaps the Mediterranean Sea. It is a robust crab of a reddish-brown colour, having an oval carapace with a characteristic "pie crust" edge and black tips to the claws. A mature adult may have a carapace width up to 25 centimetres and weigh up to 3 kilograms. C. pagurus is a nocturnal predator, targeting a range of molluscs and crustaceans. It is the subject of the largest crab fishery in Western Europe, centred on the coasts of the British Isles, with more than 60,000 tonnes caught annually.

Carcinus maenas

Carcinus maenas

Carcinus maenas is a common littoral crab. It is known by different names around the world. In the British Isles, it is generally referred to as the shore crab, or green shore crab. In North America and South Africa, it bears the name european green crab.

Liocarcinus depurator

Liocarcinus depurator

Liocarcinus depurator, sometimes called the harbour crab or sandy swimming crab, is a species of crab found in the North Sea, Atlantic Ocean, Mediterranean Sea, and Black Sea. It grows up to 50 millimetres (2.0 in) in width and 40 mm (1.6 in) long, and can be distinguished from other crabs, such as the shore crab Carcinus maenas, by the curved rows of white spots on its carapace.

Anthopleura ballii

Anthopleura ballii

Anthopleura ballii, commonly known as the red speckled anemone, is a species of sea anemone in the family Actiniidae. It is found in shallow water in the northeastern Atlantic Ocean.

Vibrio pectenicida

Vibrio pectenicida

Vibrio pectenicida is a pathogenic bacterium which attacks larvae of the scallop Pecten maximus. This bacterium does not use glucose or fructose as its carbon sources, but instead uses rhamnose and betaine. A365 is the type strain.

Fisheries and aquaculture

In 1999 the total catch reported by the United Nations Food and Agriculture Organisation was 35,411 tonnes with the two biggest catches being reported from the United Kingdom and France which landed 19,108 tonnes and 12,745 tonnes respectively.[3] It is believed that some natural stocks are showing indications of over exploitation resulting in strict enforcement of fisheries legislation and by the development of stock enhancement practices.[14] Great scallops are fished for using Newhaven scallop dredges, and less than 5% is gathered by hand by divers. In total the scallop fisheries for P. maximus and for the Queen scallop Aequipecten opercularis are one of the top five fisheries by value in United Kingdom waters. However, the use of towed great to harvest scallops causes damage to the wider ecosystem.[7]

Pecten maximus can be cultivated in aquaculture and this is reasonably advanced in France[15] and Norway.[16] Spain, France, Ireland, United Kingdom and Norway have been involved in the aquaculture of scallops; production peaked in 1998 when 512 tonnes were landed but production later recessed, with only 213 tonnes landed in 2004, having an estimated value of €852 000, equivalent to €4 per kilogramme.[6]

Pecten maximus has been found to contain domoic acid, a toxin that can cause Amnesic Shellfish Poisoning. The risk associated with scallop consumption is regarded as a significant threat to both public health and the shellfish industry.[5]

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United Kingdom

United Kingdom

The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and Northern Ireland. The United Kingdom includes the island of Great Britain, the north-eastern part of the island of Ireland, and many smaller islands within the British Isles. Northern Ireland shares a land border with the Republic of Ireland; otherwise, the United Kingdom is surrounded by the Atlantic Ocean, the North Sea, the English Channel, the Celtic Sea and the Irish Sea. The total area of the United Kingdom is 242,495 square kilometres (93,628 sq mi), with an estimated 2023 population of over 68 million people.

France

France

France, officially the French Republic, is a country located primarily in Western Europe. It also includes overseas regions and territories in the Americas and the Atlantic, Pacific and Indian Oceans, giving it one of the largest discontiguous exclusive economic zones in the world. Its metropolitan area extends from the Rhine to the Atlantic Ocean and from the Mediterranean Sea to the English Channel and the North Sea; overseas territories include French Guiana in South America, Saint Pierre and Miquelon in the North Atlantic, the French West Indies, and many islands in Oceania and the Indian Ocean. Its eighteen integral regions span a combined area of 643,801 km2 (248,573 sq mi) and had a total population of over 68 million as of January 2023. France is a unitary semi-presidential republic with its capital in Paris, the country's largest city and main cultural and commercial centre; other major urban areas include Marseille, Lyon, Toulouse, Lille, Bordeaux, and Nice.

Scuba diving

Scuba diving

Scuba diving is a mode of underwater diving whereby divers use breathing equipment that is completely independent of a surface air supply. The name "scuba", an acronym for "Self-Contained Underwater Breathing Apparatus", was coined by Christian J. Lambertsen in a patent submitted in 1952. Scuba divers carry their own source of breathing gas, usually compressed air, affording them greater independence and movement than surface-supplied divers, and more time underwater than free divers. Although the use of compressed air is common, a gas blend with a higher oxygen content, known as enriched air or nitrox, has become popular due to the reduced nitrogen intake during long and/or repetitive dives. Also, breathing gas diluted with helium may be used to reduce the likelihood and effects of nitrogen narcosis during deeper dives.

Aquaculture

Aquaculture

Aquaculture, also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants. Aquaculture involves cultivating freshwater, brackish water and saltwater populations under controlled or semi-natural conditions, and can be contrasted with commercial fishing, which is the harvesting of wild fish. Mariculture, commonly known as marine farming, refers specifically to aquaculture practiced in seawater habitats and lagoons, as opposed to freshwater aquaculture. Pisciculture is a type of aquaculture that consists of fish farming to obtain fish products as food.

Domoic acid

Domoic acid

Domoic acid (DA) is a kainic acid-type neurotoxin that causes amnesic shellfish poisoning (ASP). It is produced by algae and accumulates in shellfish, sardines, and anchovies. When sea lions, otters, cetaceans, humans, and other predators eat contaminated animals, poisoning may result. Exposure to this compound affects the brain, causing seizures, and possibly death.

Cultural significance

The oil company Shell plc derives its highly recognizable logo from this species.[17]

Pilgrims travelling to the town of Santiago de Compostella in Galicia took the shells of scallop with them, in honor of Saint James. This gave rise to the alternative name St James shell.[17]

Source: "Pecten maximus", Wikipedia, Wikimedia Foundation, (2023, February 10th), https://en.wikipedia.org/wiki/Pecten_maximus.

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References
  1. ^ a b "Pecten maximus (Linnaeus, 1758)". World Registry of Marine Species. Retrieved 1 January 2017.
  2. ^ Latchford, John W.; Beaumont, Andy R.; Wilding, Craig S. (October 1999). "Are Pecten maximus and Pecten jacobaeus different species?". Journal of the Marine Biological Association of the United Kingdom. 79 (5): 949–952. doi:10.1017/S0025315499001149. S2CID 84757441.
  3. ^ a b c d "Pecten maximus (Linnaeus, 1758)". Food and Agriculture Organization of the United Nations Department of Fisheries and Aquaculture. Retrieved 1 January 2017.
  4. ^ a b "Mollusca of the North Sea M.J. de Kluijver, S.S. Ingalsuo & R.H. de Bruyne Pecten maximus". Marine Species Identification Portal. ETI BioInformatics. Retrieved 1 January 2017.
  5. ^ a b c d e f g h "BIOTIC Species Information for Pecten maximus". BIOTIC (Biological Traits Information Catalogue). MarLIN (Marine Life Information Network). Retrieved 1 January 2016.
  6. ^ a b c A. Beaumont; T. Gjedrem (2007). Scallops - Pecten maximus and P. jacobaeus (Report). Genimpact. Retrieved 1 January 2017.
  7. ^ a b Bryce D. Beukers-Stewart; Joanne S. Beukers-Stewart (2009). Principles for the Management of Inshore Scallop Fisheries around the UK (PDF) (Report). University of York. Retrieved 1 January 2016.
  8. ^ a b Potts, Robert W. A.; Gutierrez, Alejandro P.; Penaloza, Carolina S.; Regan, Tim; Bean, Tim P.; Houston, Ross D. (2021-04-05). "Potential of genomic technologies to improve disease resistance in molluscan aquaculture". Philosophical Transactions of the Royal Society B. The Royal Society. 376 (1825): 20200168. doi:10.1098/rstb.2020.0168. ISSN 0962-8436. PMC 8059958. PMID 33813884.
  9. ^ a b Yang, Zhihui; Zhang, Lingling; Hu, Jingjie; Wang, Jing; Bao, Zhenmin; Wang, Shi (2020-04-14). "The evo‐devo of molluscs: Insights from a genomic perspective". Evolution & Development. Wiley. 22 (6): 409–424. doi:10.1111/ede.12336. ISSN 1520-541X. PMID 32291964. S2CID 215774459.
  10. ^ N. C. H. Lake; M. B. Jones; J. D. Paul (1987). "Crab predation on scallop (Pecten maximus) and its implication for scallop cultivation". Journal of the Marine Biological Association of the United Kingdom. 67 (1): 55–64. doi:10.1017/s0025315400026357. S2CID 84831172.
  11. ^ Dan Minchin (1982). "Predation on young Pecten maximus (L.) (Bivalvia), by the anemone Anthopleura ballii (Cocks)". Journal of Molluscan Studies. 49 (3): 228–231. doi:10.1093/oxfordjournals.mollus.a065717.
  12. ^ C Lambert; JL Nicolas; V Cilia; S Corre (1998). "Vibrio pectenicida sp. nov., a pathogen of scallop (Pecten maximus) larvae". International Journal of Systematic Bacteriology. 48 (2): 481–487. doi:10.1099/00207713-48-2-481. PMID 9731288.
  13. ^ Lise Thorkildsen; Christophe Lambert; Are Nylund; Thorolf Magnesen; Øivind Bergh (2005). "Bacteria associated with early life stages of the great scallop, Pecten maximus: impact on larval survival" (PDF). Aquaculture International. 13 (6): 575–592. doi:10.1007/s10499-005-9002-5. S2CID 20819269.
  14. ^ R.P. Briggs (2000). "The great scallop: an endangered species". Biologist. 47 (5): 260–264. PMID 11153135.
  15. ^ Nicole Devauchelle; Christian Mingant (1991). "Review of the reproductive physiology of the scallop, Pecten maximus, applicable to intensive aquaculture" (PDF). Aquatic Living Resources. 2: 41–51.
  16. ^ Øivind Bergh; Øivind Strand (2001). "Great scallop, Pecten maximus, research and culture strategies in Norway: a review". Aquaculture International. 9 (4): 305–317. doi:10.1023/A:1020452715518. S2CID 45170157.
  17. ^ a b "Great scallop". Ecomare. Retrieved 1 January 2016.
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