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Scallop () is a common name that encompasses various species of marine bivalve mollusks in the taxonomic
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Pectinidae, the scallops. However, the common name "scallop" is also sometimes applied to species in other closely related families within the superfamily
Pectinoidea The Pectinoidea are a superfamily of marine bivalve molluscs, including the scallops and spiny oysters.Gofas, S. (2012). Pectinoidea. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=15 ...
, which also includes the
thorny oyster ''Spondylus'' is a genus of bivalve molluscs, the only genus in the family Spondylidae.MolluscaBase (2019). MolluscaBase. Spondylus Linnaeus, 1758. Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=t ...
s. Scallops are a
cosmopolitan Cosmopolitan may refer to: Food and drink * Cosmopolitan (cocktail), also known as a "Cosmo" History * Rootless cosmopolitan, a Soviet derogatory epithet during Joseph Stalin's anti-Semitic campaign of 1949–1953 Hotels and resorts * Cosmopoli ...
family of bivalves found in all of the world's oceans, although never in fresh water. They are one of the very few groups of bivalves to be primarily "free-living", with many species capable of rapidly swimming short distances and even migrating some distance across the ocean floor. A small minority of scallop species live cemented to rocky substrates as adults, while others attach themselves to stationary or rooted objects such as seagrass at some point in their lives by means of a filament they secrete called a
byssal thread 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 (Mytili ...
. The majority of species, however, live recumbent on sandy substrates, and when they sense the presence of a predator such as a starfish, they may attempt to escape by swimming swiftly but erratically through the water using jet propulsion created by repeatedly clapping their shells together. Scallops have a well-developed nervous system, and unlike most other bivalves all scallops have a ring of numerous
simple eye Eyes are Organ (anatomy), organs of the visual system. They provide living organisms with Visual perception, vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of ...
s situated around the edge of their mantles. Many species of scallops are highly prized as a food source, and some are farmed as aquaculture. The word "scallop" is also applied to the meat of these bivalves, the adductor muscle, that is sold as seafood. The brightly coloured, symmetric, fan-shaped shells of scallops with their radiating and often fluted
ornamentation An ornament is something used for decoration. Ornament may also refer to: Decoration *Ornament (art), any purely decorative element in architecture and the decorative arts *Biological ornament, a characteristic of animals that appear to serve on ...
are valued by shell collectors, and have been used since ancient times as motifs in art, architecture, and design. Owing to their widespread distribution, scallop shells are a common sight on beaches and are often brightly coloured, making them a popular object to collect among beachcombers and vacationers. The shells also have a significant place in popular culture.


Biology


Distribution and habitat

Scallops inhabit all the oceans of the world, with the largest number of species living in the Indo-Pacific region. Most species live in relatively shallow waters from the low tide line to 100 m, while others prefer much deeper water. Although some species only live in very narrow environments, most are opportunistic and can live under a wide variety of conditions. Scallops can be found living within, upon, or under either rocks, coral, rubble,
sea grass Seagrasses are the only flowering plants which grow in marine (ocean), marine environments. There are about 60 species of fully marine seagrasses which belong to four Family (biology), families (Posidoniaceae, Zosteraceae, Hydrocharitaceae an ...
,
kelp Kelps are large brown algae seaweeds that make up the order Laminariales. There are about 30 different genera. Despite its appearance, kelp is not a plant - it is a heterokont, a completely unrelated group of organisms. Kelp grows in "underwa ...
, sand, or mud. Most scallops begin their lives as byssally attached juveniles, an ability that some retain throughout their lives while others grow into free-living adults.


Anatomy and physiology

Very little variation occurs in the internal arrangement of organs and systems within the scallop family, and what follows can be taken to apply to the anatomy of any given scallop species.


Orientation

The shell of a scallop consists of two sides or
valves A valve is a device or natural object that regulates, directs or controls the flow of a fluid (gases, liquids, fluidized solids, or slurries) by opening, closing, or partially obstructing various passageways. Valves are technically fitting ...
, a left valve and a right one, divided by a
plane of symmetry In mathematics, reflection symmetry, line symmetry, mirror symmetry, or mirror-image symmetry is symmetry with respect to a reflection. That is, a figure which does not change upon undergoing a reflection has reflectional symmetry. In 2D the ...
. Most species of scallops rest on their right valve, and consequently, this valve is often deeper and more rounded than the left (i.e., upper) valve, which in many species is actually concave. With the hinge of the two valves oriented towards the top, one side corresponds to the animal's morphological ''anterior'' or front, the other is the ''posterior'' or rear, the hinge is the ''dorsal'' or back/top region, and the bottom corresponds to the ''ventral'' or (as it were) underside/belly. However, as many scallop shells are more or less bilaterally symmetrical ("equivalved"), as well as symmetrical front/back ("equilateral"), determining which way a given animal is "facing" requires detailed information about its valves.


Valves

The model scallop shell consists of two similarly shaped valves with a straight hinge line along the top, devoid of teeth, and producing a pair of flat wings or "ears" (sometimes called "auricles", though this is also the term for two chambers in its heart) on either side of its midpoint, a feature which is unique to and apparent in all adult scallops. These ears may be of similar size and shape, or the anterior ear may be somewhat larger (the posterior ear is never larger than the anterior one, an important feature for distinguishing which valve is which). As is the case in almost all bivalves, a series of lines and/or growth rings originates at the center of the hinge, at a spot called the "
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for foo ...
" surrounded by a generally raised area called the " umbo". These growth rings increase in size downwards until they reach the curved ventral edge of the shell. The shells of most scallops are streamlined to facilitate ease of movement during swimming at some point in their lifecycles, while also providing protection from predators. Scallops with ridged valves have the advantage of the architectural strength provided by these ridges called "ribs", although the ribs are somewhat costly in weight and mass. A unique feature of the scallop family is the presence, at some point during the animal's lifecycle, of a distinctive and taxonomically important shell feature, a comb-like structure called a ctenolium located on the anterior edge of the ''right'' valve next to the valve's byssal notch. Though many scallops lose this feature as they become free-swimming adults, all scallops have a ctenolium at some point during their lives, and no other bivalve has an analogous shell feature. The ctenolium is found in modern scallops only; both putative ancestors of modern scallops, the entoliids and the Aviculopectinidae, did not possess it.


Muscular system

Like the true oysters (family
Ostreidae The Ostreidae, the true oysters, include most species of molluscs commonly consumed as oysters. Pearl oysters are not true oysters, and belong to the order Pteriida. Like scallops, true oysters have a central adductor muscle, which means the ...
), scallops have a single central adductor muscle, thus, the inside of their shells has a characteristic central scar, marking the point of attachment for this muscle. The adductor muscle of scallops is larger and more developed than those of
oyster Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not ...
s, because scallops are active swimmers; some species of scallops are known to move ''en masse'' from one area to another. In scallops, the shell shape tends to be highly regular, and is commonly used as an
archetypal The concept of an archetype (; ) appears in areas relating to behavior, historical psychology, and literary analysis. An archetype can be any of the following: # a statement, pattern of behavior, prototype, "first" form, or a main model that o ...
form of a seashell.


Digestive system

Scallops are filter feeders, and eat
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
. Unlike many other bivalves, they lack siphons. Water moves over a filtering structure, where food particles become trapped in mucus. Next, the cilia on the structure move the food toward the mouth. Then, the food is digested in the digestive gland, an organ sometimes misleadingly referred to as the "liver, " which envelops part of the oesophagus, intestine, and entire stomach. Waste is passed on through the intestine (the terminus of which, like that of many mollusks, enters and leaves the animal's heart) and exits via the anus.


Nervous system

Like all bivalves, scallops lack actual brains. Instead, their nervous system is controlled by three paired ganglia located at various points throughout their anatomy, the cerebral or cerebropleural ganglia, the pedal ganglia, and the visceral or parietovisceral ganglia. All are yellowish. The visceral ganglia are by far the largest and most extensive of the three, and occur as an almost-fused mass near the center of the animal – proportionally, these are the largest and most intricate sets of ganglia of any modern bivalve. From this, radiate all of the nerves which connect the visceral ganglia to the circumpallial nerve ring which loops around the mantle and connects to all of the scallop's tentacles and eyes. This nerve ring is so well developed that, in some species, it may be legitimately considered an additional ganglion. The visceral ganglia are also the origin of the branchial nerves which control the scallop's gills. The cerebral ganglia are the next-largest set of ganglia and lie distinct from each other a significant distance dorsal to the visceral ganglia. They are attached to the visceral ganglia by long cerebral-visceral connectives, and to each other via a cerebral commissure that extends in an arch dorsally around the esophagus. The cerebral ganglia control the scallop's mouth via the palp nerves and connect to
statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, and crustaceans. A similar structure is also found in '' Xenoturbella''. The statocyst c ...
s which help the animal sense its position in the surrounding environment. They are connected to the pedal ganglia by short cerebral-pedal connectives. The pedal ganglia, though not fused, are situated very close to each other near the midline. From the pedal ganglia, the scallop puts out pedal nerves which control the movement of, and sensation in, its small muscular foot.


Reproduction

The scallop family is unusual in that some members of the family are dioecious (males and females are separate), while others are simultaneous hermaphrodites (both sexes in the same individual), and a few are protoandrous hermaphrodites (males when young then switching to female). Red
roe Roe ( ) or hard roe is the fully ripe internal egg masses in the ovaries, or the released external egg masses, of fish and certain marine animals such as shrimp, scallop, sea urchins and squid. As a seafood, roe is used both as a cooked in ...
is that of a female, and white, is that of a male. Spermatozoa and ova are released freely into the water during mating season and fertilized ova sink to the bottom. After several weeks, the immature scallops hatch and the larvae, miniature transparent versions of the adults called "spat", drift in the plankton until settling to the bottom again (an event called spatfall) to grow, usually attaching by means of byssal threads. Some scallops, such as the Atlantic bay scallop ''Argopecten irradians'', are short-lived, while others can live 20 years or more. Age can often be inferred from annuli, the concentric rings of their shells.


Eyes

Scallops have a large number (up to 200) of small (about 1 mm) eyes arranged along the edge of their mantles. These eyes represent a particular innovation among molluscs, relying on a concave, parabolic mirror of
guanine Guanine () ( symbol G or Gua) is one of the four main nucleobases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine (uracil in RNA). In DNA, guanine is paired with cytosine. The guanine nucleoside is c ...
crystals to focus and retro-reflect light instead of a lens as found in many other eye types. Additionally, their eyes possess a double-layered retina, the outer retina responding most strongly to light and the inner to abrupt darkness. While these eyes are unable to resolve shapes with high fidelity, the combined sensitivity of both retinas to light entering the eye and light retro-reflected from the mirror grants scallops exceptional contrast definition, as well as the ability to detect changing patterns of light and motion. Scallops primarily rely on their eyes as an 'early-warning' threat detection system, scanning around them for movement and shadows which could potentially indicate predators. Additionally, some scallops alter their swimming or feeding behaviour based on the turbidity or clarity of the water, by detecting the movement of particulate matter in the water column.


Adductor muscles

Scallops possess fast ( striated) and slow (
smooth Smooth may refer to: Mathematics * Smooth function, a function that is infinitely differentiable; used in calculus and topology * Smooth manifold, a differentiable manifold for which all the transition maps are smooth functions * Smooth algebrai ...
) adductor muscles, which have different structures and contractile properties. These muscles lie closely apposed to one another but are divided by a connective tissue sheet. The striated adductor muscle contracts very quickly for swimming, whereas the smooth catch adductor muscle lacks striations, and contracts for long periods, keeping shells closed with little expenditure of energy. Material was copied from this source, which is available under
Creative Commons Attribution 4.0 International License


Locomotion

Scallops are mostly free-living and active, unlike the vast majority of bivalves, which are mostly slow-moving and infaunal. All scallops are thought to start out with a
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 (Mytil ...
, which attaches them to some form of substrate such as
eelgrass Eelgrass is a common name for several plants and may refer to: * ''Zostera'', marine eelgrass * ''Vallisneria'', freshwater eelgrass {{Short pages monitor The scallop shell is the traditional emblem of
St James the Great James the Great, also known as James, son of Zebedee, Saint James the Great, Saint James the Greater, Saint James the Elder, or Saint Jacob (Aramaic ܝܥܩܘܒ ܒܪ ܙܒܕܝ, Arabic يعقوب, Hebrew בן זבדי , '' Yaʿăqōḇ'', Latin '' ...
and is popular with pilgrims returning from the Way of St James ( Camino de Santiago) and the apostle's shrine at
Santiago de Compostela Santiago de Compostela is the capital of the autonomous community of Galicia, in northwestern Spain. The city has its origin in the shrine of Saint James the Great, now the Cathedral of Santiago de Compostela, as the destination of the Way of S ...
in Galicia,
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
. Medieval Christians would collect a scallop shell while at Compostela as evidence of having made the journey. The association of Saint James with the scallop can most likely be traced to the legend that the apostle once rescued a knight covered in scallops. An alternative version of the legend holds that while St. James' remains were being transported to Galicia (Spain) from
Jerusalem Jerusalem (; he, יְרוּשָׁלַיִם ; ar, القُدس ) (combining the Biblical and common usage Arabic names); grc, Ἱερουσαλήμ/Ἰεροσόλυμα, Hierousalḗm/Hierosóluma; hy, Երուսաղեմ, Erusałēm. i ...
, the disciples witnessed a bridegroom on horseback fall into the water and emerge covered in the shells. Indeed in French, the mollusc itself – as well as a popular preparation of it in cream sauce – is called . In German they are – literally "James's shellfish". Curiously the Linnaean name ''
Pecten jacobeus ''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 ''Pecten jacobaeus'' usually reaches a length of about , but ...
'' is given to the Mediterranean scallop, while the scallop endemic to Galicia is called ''Pecten maximus'' due to its bigger size. The scallop shell is represented in the decoration of churches named after St. James, such as in
St James' Church, Sydney St James' Church, commonly known as St James', King Street, is an Australian heritage-listed Anglican parish church located at 173 King Street, in the Sydney central business district in New South Wales. Consecrated in February 1824 and named ...
, where it appears in a number of places, including in the mosaics on the floor of the
chancel In church architecture, the chancel is the space around the altar, including the choir and the sanctuary (sometimes called the presbytery), at the liturgical east end of a traditional Christian church building. It may terminate in an apse. Ov ...
. When referring to St James, a scallop shell valve is displayed with its convex outer surface showing. In contrast, when the shell refers to the goddess Venus (see below), it is displayed with its concave interior surface showing.


Shell of Saint Augustine

Saint Augustine Augustine of Hippo ( , ; la, Aurelius Augustinus Hipponensis; 13 November 354 – 28 August 430), also known as Saint Augustine, was a theologian and philosopher of Berber origin and the bishop of Hippo Regius in Numidia, Roman North Afr ...
is said to have been walking along the seashore, meditating on the unfathomable mystery of the Holy Trinity. A boy was using a shell to pour seawater into a little hole. When Augustine asked what he was doing, he replied, "I am emptying the sea into this hole." Thus did
Augustine Augustine of Hippo ( , ; la, Aurelius Augustinus Hipponensis; 13 November 354 – 28 August 430), also known as Saint Augustine, was a theologian and philosopher of Berber origin and the bishop of Hippo Regius in Numidia, Roman North A ...
understand that man would never penetrate to the depths of the mystery of God. This symbolic meaning was taken up by
Joseph Ratzinger Pope Benedict XVI ( la, Benedictus XVI; it, Benedetto XVI; german: link=no, Benedikt XVI.; born Joseph Aloisius Ratzinger, , on 16 April 1927) is a retired prelate of the Catholic church who served as the head of the Church and the soverei ...
in his coat of arms as Archbishop of Munich, and also retained by him when elected Pope
Benedict XVI Pope Benedict XVI ( la, Benedictus XVI; it, Benedetto XVI; german: link=no, Benedikt XVI.; born Joseph Aloisius Ratzinger, , on 16 April 1927) is a retired prelate of the Catholic church who served as the head of the Church and the soverei ...
. While a doctoral candidate in 1953, Ratzinger wrote his dissertation on ''The People of God and the House of God in Augustine's Teaching'', and the shell, therefore, has a personal connection with the thought of Saint Augustine.


Badge

The scallop shell symbol found its way into heraldry as a badge of those who had been on the pilgrimage to Compostela, although later, it became a symbol of pilgrimage in general.
Winston Churchill Sir Winston Leonard Spencer Churchill (30 November 187424 January 1965) was a British statesman, soldier, and writer who served as Prime Minister of the United Kingdom twice, from 1940 to 1945 during the Second World War, and again from ...
and Diana, Princess of Wales' family, the
Spencer family The Spencer family is an aristocratic family in the United Kingdom. From the 16th century, its members have held numerous titles including the dukedom of Marlborough, the earldoms of Sunderland and Spencer, and the Churchill barony. Two prom ...
coat of arms A coat of arms is a heraldic visual design on an escutcheon (i.e., shield), surcoat, or tabard (the latter two being outer garments). The coat of arms on an escutcheon forms the central element of the full heraldic achievement, which in its ...
includes a scallop, as well as both of Diana's sons Prince William, Duke of Cambridge and
Prince Harry Prince Harry, Duke of Sussex, (Henry Charles Albert David; born 15 September 1984) is a member of the British royal family. He is the younger son of Charles III and his first wife Diana, Princess of Wales. He is fifth in the line of succ ...
's personal coats of arms; also Pope
Benedict XVI Pope Benedict XVI ( la, Benedictus XVI; it, Benedetto XVI; german: link=no, Benedikt XVI.; born Joseph Aloisius Ratzinger, , on 16 April 1927) is a retired prelate of the Catholic church who served as the head of the Church and the soverei ...
's personal coat of arms includes a scallop; another example is the surname
Wilmot Wilmot may refer to: Places Australia *Division of Wilmot, an abolished Australian Electoral Division in Tasmania * Wilmot, Tasmania, a locality in the North-West Region Canada *Wilmot, Nova Scotia, an unincorporated rural community and former t ...
and also John Wesley's (which as a result the scallop shell is used as an emblem of
Methodism Methodism, also called the Methodist movement, is a group of historically related denominations of Protestant Christianity whose origins, doctrine and practice derive from the life and teachings of John Wesley. George Whitefield and John's br ...
). However, charges in heraldry do not always have an unvarying symbolic meaning, and there are cases of arms in which no family member went on a pilgrimage, and the occurrence of the scallop is simply a pun on the name of the
armiger In heraldry, an armiger is a person entitled to use a heraldic achievement (e.g., bear arms, an "armour-bearer") either by hereditary right, grant, matriculation, or assumption of arms. Such a person is said to be armigerous. A family or a cl ...
(as in the case of Jacques Coeur), or for other reasons. In 1988, the State of New York in the US chose the bay scallop (''Argopecten irradians'') as its state shell.


Fertility symbol

Throughout antiquity, scallops and other hinged shells have symbolized the feminine principle. Outwardly, the shell can symbolize the protective and nurturing principle, and inwardly, the "life-force slumbering within the Earth", an emblem of the vulva. Many paintings of
Venus Venus is the second planet from the Sun. It is sometimes called Earth's "sister" or "twin" planet as it is almost as large and has a similar composition. As an interior planet to Earth, Venus (like Mercury) appears in Earth's sky never f ...
, the Roman goddess of love and fertility, included a scallop shell in the painting to identify her. This is evident in Botticelli's classically inspired 15th century painting ''
The Birth of Venus ''The Birth of Venus'' ( it, Nascita di Venere ) is a painting by the Italian artist Sandro Botticelli, probably executed in the mid 1480s. It depicts the goddess Venus arriving at the shore after her birth, when she had emerged from the sea ...
''. One legend of the Way of St. James holds that the route was seen as a fertility pilgrimage, undertaken when a young couple desired to bear offspring. The scallop shell is believed to have originally been carried by pagans as a symbol of fertility.


Other interpretations

Alternatively, the scallop resembles the setting sun, which was the focus of the pre-Christian Celtic rituals of the area. To wit, the pre-Christian roots of the ''Way of St. James'' was a Celtic death journey westwards towards the setting sun, terminating at the ''End of the World'' (''Finisterra'') on the "Coast of Death" (Costa da Morte) and the "Sea of Darkness" (i.e., the Abyss of Death, the ''Mare Tenebrosum'', Latin for the Atlantic Ocean, itself named after the ''Dying Civilization'' of Atlantis).


Contemporary art

The beach at Aldeburgh, Suffolk, England, features Maggi Hambling's steel sculpture, ''The Scallop'', erected in 2003 as a memorial to the composer Benjamin Britten, who had a long association with the town.


See also

*


Explanatory notes


Citations


General bibliography

* * * * * * * * * * * * * * * * * * * * * * *


External links


Mollusca - Bivalvia - Pectinidae
at Natural History Museum Rotterdam – photos of Pectinidae shells {{Taxonbar, from=Q1311395 Pectinidae, Bivalve families Commercial molluscs Extant Middle Triassic first appearances Heraldic charges Mollusc common names