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In most biological
nomenclature Nomenclature (, ) is a system of names or terms, or the rules for forming these terms in a particular field of arts or sciences. The principles of naming vary from the relatively informal naming conventions, conventions of everyday speech to the i ...
, a scale ( grc, λεπίς, lepís; la, squāma) is a small rigid
plate Plate may refer to: Cooking * Plate (dishware), a broad, mainly flat vessel commonly used to serve food * Plates, tableware, dishes or dishware used for setting a table, serving food and dining * Plate, the content of such a plate (for example: ...
that grows out of an
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
's
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other cuticle, animal coverings, such as the arthropod exoskeleton, have diffe ...
to provide protection. In
lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
n (
butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
and
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
) species, scales are plates on the surface of the
insect wing Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindwin ...
, and provide coloration. Scales are quite common and have evolved multiple times through
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
, with varying structure and function. Scales are generally classified as part of an organism's
integumentary system The integumentary system is the set of organs forming the outermost layer of an animal's body. It comprises the skin and its appendages, which act as a physical barrier between the external environment and the internal environment that it serves ...
. There are various types of scales according to shape and to
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
of animal.


Fish scales

File:Ganoid scales.png, Ganoid scales on a
carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
fish ''
Amblypterus ''Amblypterus'' (from el, ἀμβλύς , 'blunt' and el, πτερόν 'wing' or 'fin') is an extinct genus of ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise ...
striatus'' File:Denticules cutanés du requin citron Negaprion brevirostris vus au microscope électronique à balayage.jpg, Placoid scales on a
lemon shark The lemon shark (''Negaprion brevirostris'') is a species of shark from the family Carcharhinidae and is classified as a Vulnerable species by the International Union for the Conservation of Nature. Lemon sharks can grow to in length. They are o ...
(''Negaprion brevirostris'') File:RutilusRutilusScalesLateralLine.JPG, Cycloid scales on a common roach (''Rutilus rutilus'')
Fish scales are dermally derived, specifically in the
mesoderm The mesoderm is the middle layer of the three germ layers that develops during gastrulation in the very early development of the embryo of most animals. The outer layer is the ectoderm, and the inner layer is the endoderm.Langman's Medical E ...
. This fact distinguishes them from reptile scales paleontologically. Genetically, the same genes involved in tooth and hair development in mammals are also involved in scale development.


Cosmoid scales

True cosmoid scales can only be found on the
Sarcopterygian Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includi ...
s. The inner layer of the scale is made of
lamellar A ''lamella'' (plural ''lamellae'') is a small plate or flake, from the Latin, and may also be used to refer to collections of fine sheets of material held adjacent to one another, in a gill-shaped structure, often with fluid in between though s ...
bone. On top of this lies a layer of spongy or
vascular The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away f ...
bone and then a layer of
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
-like material called
cosmine Cosmine is a spongy, bony material that makes up the dentine-like layers in the scales of the lobe-finned fishes of the class Sarcopterygii. Fish scales that include layers of cosmine are known as cosmoid scales. Description As traditionally des ...
. The upper surface is
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
. The
coelacanth The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
has modified cosmoid scales that lack cosmine and are thinner than true cosmoid scales.


Ganoid scales

Ganoid scales can be found on
gar Gars are members of the family Lepisosteidae, which are the only surviving members of the Ginglymodi, an ancient holosteian group of ray-finned fish, which first appeared during the Triassic, over 240 million years ago. Gars comprise seven livin ...
s (family
Lepisosteidae Gars are members of the family Lepisosteidae, which are the only surviving members of the Ginglymodi, an ancient holosteian group of ray-finned fish, which first appeared during the Triassic, over 240 million years ago. Gars comprise seven livin ...
),
bichir Bichirs and the reedfish comprise Polypteridae , a family of archaic ray-finned fishes and the only family in the order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes. West Sussex, UK: Blackwell Pub ...
s, and reedfishes (family
Polypteridae Bichirs and the reedfish comprise Polypteridae , a family of archaic ray-finned fishes and the only family in the order Polypteriformes .Helfman GS, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes. West Sussex, UK: Blackwell Pub ...
). Ganoid scales are similar to cosmoid scales, but a layer of
ganoin Ganoine or ''ganoin'' is a glassy, often multi-layered mineralized tissue that covers the scales, cranial bones and fin rays in some non-teleost ray-finned fishes, such as gars, bichirs, and coelacanths. It is composed of rod-like, pseudoprismat ...
lies over the cosmine layer and under the enamel. Ganoin scales are diamond shaped, shiny, and hard. Within the ganoin are
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 called ...
compounds, iridescent derivatives of guanine found in a DNA molecule. The iridescent property of these chemicals provide the ganoin its shine.


Placoid scales

Placoid scale A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as ...
s are found on cartilaginous fish including
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s and
stingray Stingrays are a group of sea rays, which are cartilaginous fish related to sharks. They are classified in the suborder Myliobatoidei of the order Myliobatiformes and consist of eight families: Hexatrygonidae (sixgill stingray), Plesiobatidae ( ...
s. These scales, also called denticles, are similar in structure to
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
, and have one median spine and two lateral spines. The modern jawed fish ancestors, the jawless
ostracoderm Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) an ...
s and later jawed placoderms, may have had scales with the properties of both placoid and ganoid scales.


Leptoid scales

Leptoid scales are found on higher-order bony fish. As they grow they add concentric layers. They are arranged so as to overlap in a head-to-tail direction, like roof tiles, allowing a smoother flow of water over the body and therefore reducing drag. They come in two forms: *Cycloid scales have a smooth outer edge, and are most common on fish with soft fin rays, such as
salmon Salmon () is the common name for several list of commercially important fish species, commercially important species of euryhaline ray-finned fish from the family (biology), family Salmonidae, which are native to tributary, tributaries of the ...
and
carp Carp are various species of oily freshwater fish from the family Cyprinidae, a very large group of fish native to Europe and Asia. While carp is consumed in many parts of the world, they are generally considered an invasive species in parts of ...
. *Ctenoid scales have a toothed outer edge, and are usually found on fish with spiny fin rays, such as bass and
crappie Crappies () are two species of North American freshwater fish of the genus ''Pomoxis'' in the family Centrarchidae (sunfishes). Both species of crappies are popular game fish among recreational anglers. Etymology The genus name ''Pomoxis'' ...
.


Reptilian scales

Reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
scale types include: cycloid, granular (which appear bumpy), and keeled (which have a center ridge). Scales usually vary in size, the stouter, larger scales cover parts that are often exposed to physical stress (usually the feet, tail and head), while scales are small around the joints for flexibility. Most snakes have extra broad scales on the belly, each scale covering the belly from side to side. The scales of all reptiles have an epidermal component (what one sees on the surface), but many reptiles, such as
crocodilian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
s and turtles, have
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...
s underlying the epidermal scale. Such scales are more properly termed
scute A scute or scutum (Latin: ''scutum''; plural: ''scuta'' "shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of birds. The term is also used to describe the anterior po ...
s. Snakes,
tuatara Tuatara (''Sphenodon punctatus'') are reptiles endemic to New Zealand. Despite their close resemblance to lizards, they are part of a distinct lineage, the order Rhynchocephalia. The name ''tuatara'' is derived from the Māori language and m ...
s and many lizards lack osteoderms. All reptilian scales have a dermal papilla underlying the epidermal part, and it is there that the osteoderms, if present, would be formed.


Avian scales

Birds' scales are found mainly on the toes and metatarsus, but may be found further up on the ankle in some birds. The scales and scutes of birds were thought to be homologous to those of reptiles, but are now agreed to have evolved independently, being degenerate feathers.


Mammalian scales

An example of a scaled mammal is the
pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (, from Ancient Greek ϕολιδωτός – "clad in scales"). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smut ...
. Its scales are made of keratin and are used for protection, similar to an
armadillo Armadillos (meaning "little armored ones" in Spanish) are New World placental mammals in the order Cingulata. The Chlamyphoridae and Dasypodidae are the only surviving families in the order, which is part of the superorder Xenarthra, along wi ...
's armor. They have been convergently evolved, being unrelated to mammals' distant reptile-like ancestors (since
therapsids Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented mor ...
lost scales), except that they use a similar gene. On the other hand, the
musky rat-kangaroo The musky rat-kangaroo (''Hypsiprymnodon moschatus'') is a small marsupial found only in the rainforests of northeastern Australia. First described in the later 19th century, the only other species are known from fossil specimens. They are simil ...
has scales on its feet and tail. The precise nature of its purported scales has not been studied in detail, but they appear to be structurally different from pangolin scales.
Anomalure The Anomaluridae are a family of rodents found in central Africa. They are known as anomalures or scaly-tailed squirrels. The six extant species are classified into two genera. All anomalurids have membranes between their front and hind legs li ...
s also have scales on their tail undersides. Foot pad epidermal tissues in most mammal species have been compared to the scales of other vertebrates. They are likely derived from cornification processes or stunted fur much like avian reticulae are derived from stunted feathers.


Arthropod scales

Butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
and
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s - the order
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
(
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
"scale-winged") - have membranous wings covered in delicate, powdery scales, which are modified
setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
. Each scale consists of a series of tiny stacked platelets of organic material, and butterflies tend to have the scales broad and flattened, while moths tend to have the scales narrower and more hair like. Scales are usually
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...
ed, but some types of scales are iridescent, without pigments; because the thickness of the platelets is on the same order as the
wavelength In physics, the wavelength is the spatial period of a periodic wave—the distance over which the wave's shape repeats. It is the distance between consecutive corresponding points of the same phase on the wave, such as two adjacent crests, tro ...
of
visible light Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 te ...
the plates lead to structural coloration and
iridescence Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear to gradually change color as the angle of view or the angle of illumination changes. Examples of iridescence include soap bubbles, feathers, butterfl ...
through the physical phenomenon described as
thin-film optics Thin-film optics is the branch of optics that deals with very thin structured layers of different materials. In order to exhibit thin-film optics, the thickness of the layers of material must be similar to the coherence length; for visible l ...
. The most common color produced in this fashion is
blue Blue is one of the three primary colours in the RYB colour model (traditional colour theory), as well as in the RGB (additive) colour model. It lies between violet and cyan on the spectrum of visible light. The eye perceives blue when obs ...
, such as in the ''
Morpho The morpho butterflies comprise many species of Neotropical butterfly under the genus ''Morpho''. This genus includes more than 29 accepted species and 147 accepted subspecies, found mostly in South America, Mexico, and Central America. ''Morph ...
'' butterflies. Some types of
spider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s also have scales. Spider scales are flattened setae that overlay the surface of the
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
. They come in a wide variety of shapes, sizes, and colors. At least 13 different spider families are known to possess cuticular scales, although they have only been well described for
jumping spider Jumping spiders are a group of spiders that constitute the family (biology), family Salticidae. As of 2019, this family contained over 600 described genera and over 6,000 described species, making it the largest family of spiders at 13% of all sp ...
s (Salticidae) and
lynx spider Lynx spider (Oxyopidae) is a family of araneomorph spiders first described by Tamerlan Thorell in 1870. Most species make little use of webs, instead spending their lives as hunting spiders on plants. Many species frequent flowers in particular, ...
s (Oxyopidae).


See also

*
Scale (insect anatomy) Scales are present on the bodies of various insects. A notable example are Lepidoptera, the insect order comprising moths and butterflies, which has scales on their wings and also on the head, parts of the thorax and abdomen as well as parts of the ...
*
Armour (anatomy) Armour or armor in animals is external or superficial protection against attack by predators, formed as part of the body (rather than the behavioural use of protective external objects), usually through the hardening of body tissues, outgrowths o ...
*
Psoriasis Psoriasis is a long-lasting, noncontagious autoimmune disease characterized by raised areas of abnormal skin. These areas are red, pink, or purple, dry, itchy, and scaly. Psoriasis varies in severity from small, localized patches to complete ...
: a long-lasting autoimmune disease characterized by patches of thin pieces of hard skin like scale.


References


Further reading

* {{refend Animal anatomy Marine biology Integumentary system