Fish scale - Knowledge (XXG)

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turbulent vortices became trapped between the denticles, creating a ‘cushion like’ barrier against the laminar flow. This same type of experiment was performed by another research group which implemented more variation in their biomimetic sample. The second group arrived at the same conclusion as the first. However, because their experiment contained more variation within the samples they were able to achieve a high degree of experimental accuracy. In conclusion, they stated that more practical shapes were more durable than ones with intricate ridge-lines. The practical shapes were low profile and contained trapezoidal or semi-circular trough-like cross sections, and were less effective but nonetheless reduced drag by 6 or 7%.

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pushes the vortex further up from the surface, interacting only with the riblet tips, not causing any high-velocity flow in the valleys. Since this high-velocity flow now only interacts with the riblet-tip, which is a very small surface area, the momentum transfer which causes drag is now much lower than before, thereby effectively reducing drag. Also, this reduces the cross-stream velocity fluctuations, which aids in momentum transfer too.

1383:, which lives in shallower water, the mirrors must reflect a mixture of wavelengths, and the fish accordingly has crystal stacks with a range of different spacings. A further complication for fish with bodies that are rounded in cross-section is that the mirrors would be ineffective if laid flat on the skin, as they would fail to reflect horizontally. The overall mirror effect is achieved with many small reflectors, all oriented vertically. 2428: 2334: 1182: 5581: 1214: 1198: 1166: 2012: 1074: 545: 2341: 1106: 6573: 6555: 1129: 1996: 1601: 1323: 209: 1609: 792: 567:. (a) shows the outer surface of four of the scales, and (b) shows the inner surface of two of the scales. Each of the rhomboidal-shaped ganoid scales of Amblypterus has a ridge on the inner surface which is produced at one end into a projecting peg which fits into a notch in the next scale, similar to the manner in which tiles are pegged together on the roof of a house. 6585: 572: 1490: 40: 1339: 280: 2491:, a fish is kosher if the scales can be removed without tearing its skin. Thus carp and salmon are kosher, whereas a shark, whose scales are microscopic, a sturgeon, whose scutes can not be easily removed without cutting them out of the body, are all not kosher. Other non-kosher fish include catfish, eels, 1925:

are composed of mineralized and non-mineralized collagen in different regions. The formation of these occurs through the entry of the surface cells of the mesenchyme into the matrix, the latter is composed of collagen fibers and is located around the vascular capillaries, thus giving rise to vascular

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are similar to scales and serve the same function. Unlike the scales of fish, which are formed from the epidermis, scutes are formed in the lower vascular layer of the skin and the epidermal element is only the top surface. Forming in the living dermis, the scutes produce a horny outer layer, that is

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to supply the denticle with mucus. Denticles contain riblet structures that protrude from the surface of the scale; under a microscope this riblet can look like a hook or ridges coming out of the scale. The overall shape of the protrusion from the denticle is dependent on the type of shark and can be

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forms. One morph has its jaw twisted to the left, allowing it to eat scales more readily on its victim's right flank. The other morph has its jaw twisted to the right, which makes it easier to eat scales on its victim's left flank. The relative abundance of the two morphs in populations is regulated

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of ray-finned fishes). The outer part of these scales fan out with bony ridges while the inner part is criss-crossed with fibrous connective tissue. Leptoid scales are thinner and more translucent than other types of scales, and lack the hardened enamel-like or dentine layers. Unlike ganoid scales,

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upper surface and an aspidine base. Its growing base is made of cell-free bone, which sometimes developed anchorage structures to fix it in the side of the fish. Beyond that, there appear to be five types of bone growth, which may represent five natural groupings within the thelodonts—or a spectrum

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spaced about ¼ of a wavelength apart to interfere constructively and achieve nearly 100 per cent reflection. In the deep waters that the hatchetfish lives in, only blue light with a wavelength of 500 nanometres percolates down and needs to be reflected, so mirrors 125 nanometres apart provide good

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traits is prone to errors. Nonetheless, a framework comprising three groups has been proposed based upon scale morphology and histology. Comparisons to modern shark species have shown that thelodont scales were functionally similar to those of modern cartilaginous fish, and likewise has allowed an

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However, using scale morphology alone to distinguish species has some pitfalls. Within each organism, scale shape varies hugely according to body area, with intermediate forms appearing between different areas—and to make matters worse, scale morphology may not even be constant within one area. To

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The riblets impede the cross-stream translation of the streamwise vortices in the viscous sublayer. The mechanism is complex and not yet understood fully. Basically, the riblets inhibit the vortex formation near the surface because the vortex cannot fit in the valleys formed by the riblets. This

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reacted. Both the geometry of the denticles and their arrangement have a profound effect on the aerodynamic response of the aerofoils. Out of both the low and high-profile samples tested, the low-profile vortex generators outperformed the current smooth wing structures by 323%. This increase in

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Denticles also provide drag reduction on objects where the main form of drag is caused by turbulent flow at the surface. A large portion of the total drag on long objects with relatively flat sides usually comes from turbulence at the wall, so riblets will have an appreciable effect. Along with

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network surrounding their body. The corset works as an outer skeleton, providing attachment for their swimming muscles and thus saving energy. Depending on the position of these placoid scales on the body, they can be flexible and can be passively erected, allowing them to change their angle of

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samples of shark denticles with a crescent like microstructure were tested in a water tank using a traction table as a slide. The experiment showed that the surface with denticles experienced a 10% drag reduction overall versus the smooth sample. The reason for this drag reduction was that the

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Differentiation or late metamorphosis: This differentiation is generated by two different forms according to the type of scale being formed. The formation of elasmoid scales (cycloids and ctenoids) occurs through the formation of a space between the matrix of the epidermal papilla. This space

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The general anatomy of the scales varies, but all of them can be divided into three parts: the crown, the neck and the base. The scale pliability is related to the size of the base of the scale. The scales with higher flexibility have a smaller base, and thus are less rigidly attached to the

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has been done on shark denticles with a wide range of design variations such as low and high-profile vortex generators. Through this method, the most thorough characterization has been completed for symmetrical two-dimensional riblets with sawtooth, scalloped and blade cross sections. These

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marine applications, the aerospace industry can benefit greatly from these biomimetic designs. Other applications include pipes, where they score the insides to a riblet-like roughness and have discovered a 5% drag reduction, and a few percent reduction is claimed with competitive swimwear.

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thickens at a declining rate and then abruptly undergoes a breakdown into turbulent vortices before finally collapsing. This system is completely self-regulating and mediates the growth and decay cycle; the vortices accumulate during the growth period and are abruptly liquidated into

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Analyzing the three components of the scale it can be concluded that the base of the denticle does not come into contact with any portion of the fluid flow. The crown and the neck of the denticles however play a key role and are responsible for creating the turbulent vortices and

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generally described with two appearances. The first is a scale in which ridges are placed laterally down the shark and parallel with the flow of the water. The second form is a smooth scale with what looks like a hooked riblet curling out of the surface aiming towards the

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have thinner, more hidden spines than porcupine fish, which become visible only when the fish puffs up. Unlike the porcupine fish, these spines are not modified scales, but develop under the control of the same network of genes that produce feathers and hairs in other

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ranging between the end members meta- (or ortho-) dentine and mesodentine tissues. Each of the five scale morphs appears to resemble the scales of more derived groupings of fish, suggesting that thelodont groups may have been stem groups to succeeding clades of fish.

1740:(PDMS) for creating a mold. Usually the process involves taking a flat piece of shark skin, covering it with the PDMS to form a mold and pouring PDMS into that mold again to get a shark skin replica. This method has been used to create a biomimetic surface which has 1350:

Many teleost fish are covered with highly reflective scales which function as small mirrors and give the appearance of silvered glass. Reflection through silvering is widespread or dominant in fish of the open sea, especially those that live in the top 100 metres. A

2483:, it must have scales and fins. The definition of "scale" differs from the definitions presented in biology, in that the scales of a kosher fish must be visible to the eye, and can be easily removed from the skin either by hand or scaling knife. According to the 1629:. When the laminar flow increases around the fish the pressure drag does as well. Frictional drag is a result of the interaction between the fluid against the shark's skin and can vary depending on how the boundary layer changes against the surface of the fish. 1862:. This process depends on whether the fish is cartilaginous or bony. For cartilaginous fish the structuring originates through the formation of two layers. The first is superficial and wide and the second is thin and compact. These two layers are separated by 584:(diamond-shaped) and connected by peg-and-socket joints. They are usually thick and fit together more like a jigsaw rather than overlapping like other scales. In this way, ganoid scales are nearly impenetrable and are excellent protection against predation. 1562:

attack. These scales also have riblets which are aligned in the direction of flow, these riblets reduce the drag force acting on the shark skin by pushing the vortex further away from the skin surface, inhibiting any high-velocity cross-stream flow.

2541:. Scale eating behaviour usually evolves because of lack of food and extreme environmental conditions. The eating of scales and the skin surrounding the scales provides protein rich nutrients that may not be available elsewhere in the niche. 1624:

is created from the pressure difference between the anterior and posterior sides of the shark due to the amount of volume that is pushed past the shark to propel itself forward. This type of drag is also directly proportional to the

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biomimetic models were designed and analyzed to see the effects of applying the denticle-like structures to the wings of various airplanes. During the simulation, it was noted that the sample altered how the low and high

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Dean, Brian & Bhushan, Bharat. (2010). Shark-Skin Surfaces for Fluid-Drag Reduction in Turbulent Flow: A Review. Philosophical transactions. Series A, Mathematical, physical, and engineering sciences. 368. 4775-806.

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Shark skin is almost entirely covered by small placoid scales. The scales are supported by spines, which feel rough when stroked in a backward direction, but when flattened by the forward movement of water, create tiny

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Motta, Philip; Habegger, Maria Laura; Lang, Amy; Hueter, Robert; Davis, Jessica (1 October 2012). "Scale morphology and flexibility in the shortfin mako Isurus oxyrinchus and the blacktip shark Carcharhinus limbatus".

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effect can be achieved by silvering to make an animal's body highly reflective. At medium depths at sea, light comes from above, so a mirror oriented vertically makes animals such as fish invisible from the side.

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declares that, "of all that are in the waters... in the seas, and in the rivers" those that do not have both fins and scales "shall be an abomination unto you" and may not be eaten. This eliminates all aquatic

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Leptoid scales overlap in a head-to-tail configuration, like roof tiles, making them more flexible than cosmoid and ganoid scales. This arrangement allows a smoother flow of water over the body, and reduces

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Sadovy, Y.; Randall, J. E.; Rasotto, Maria B. (May 2005). "Skin structure in six dragonet species (Gobiesociformes; Callionymidae): Interspecific differences in glandular cell types and mucus secretion".

1711:. Dermal denticles are an extremely promising area of research for this type of application due to the fact that sharks are among the only fish without build up or growth on their scales. Studies by the 1645:

of hairpin vortices lifting off the wall. Lifting vortices are what push the boundary layer out and away from the surface of the shark which results in reducing the overall drag experienced by the fish.

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de Vrieze, E.; van Kessel, M. A. H. J.; Peters, H. M.; Spanings, F. A. T.; Flik, G.; Metz, J. R. (1 February 2014). "Prednisolone induces osteoporosis-like phenotype in regenerating zebrafish scales".

4586: 2193: 1842:, it takes 30 days after fertilization before the different layers needed to start forming the scales have differentiated and become organized. For this it is necessary that consolidation of the 3479:

Pouyaud, L.; Sudarto, Guy G. Teugels (2003). "The different colour varieties of the Asian arowana Scleropages formosus (Osteoglossidae) are distinct species: morphologic and genetic evidences".

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and surfaces based on the structure of aquatic organisms, including sharks. Such applications intend to enable more efficient movement through fluid mediums such as air, water, and oil.

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a projecting, modified (rough and strongly ridged) scale, usually associated with the lateral line, or on the caudal peduncle forming caudal keels, or along the ventral profile.

2405:, a protective band of thickened and enlarged scales in the shoulder region. Over most of their body tuna have scales so small that to casual inspection they seems scaleless. 1913:

contains collagen fibers. Around this space elasmoblasts differentiate and are responsible for generating the necessary material for the formation of the scale. Subsequently,

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Fürstner, Reiner; Barthlott, Wilhelm; Neinhuis, Christoph; Walzel, Peter (1 February 2005). "Wetting and Self-Cleaning Properties of Artificial Superhydrophobic Surfaces".

2745: 2286:, but they do not have regular scales. They rely on camouflage to avoid the attention of predators, while their loose skin makes it difficult for predators to grab them. 884: 6609: 5426: 2394:

Filefish have rough non-overlapping scales with small spikes, which is why they are called filefish. Some filefish appear scaleless because their scales are so small.

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In sturgeons, the scales are greatly enlarged into armour plates along the sides and back, while in the bowfin the scales are greatly reduced in thickness to resemble

630: 2411: 2362: 590: 2537:. However, the energy expended to make a strike versus the amount of scales consumed per strike puts a limit on the size of lepidophagous fish, and they are usually 2174: 266:

The morphology and histology of thelodonts provides the main tool for quantifying their diversity and distinguishing between species, although ultimately using such

5349:"Evolution of a unique predatory feeding apparatus: functional anatomy, development and a genetic locus for jaw laterality in Lake Tanganyika scale-eating cichlids" 4772:

Sorenson, L.; Santini, F.; Carnevale, G.; Alfaro, M.E. (2013). "A multi-locus timetree of surgeonfishes (Acanthuridae, Percomorpha), with revised family taxonomy".

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performance is due to a separation bubble in the denticle's wake and stream-wise vortices that replenish momentum lost in the boundary layer due to skin friction.

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found near the skin's surface. Because denticles come in so many different shapes and sizes, it can be expected that not all shapes will produce the same type of

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in a modified form (see elasmoid scales, below). They were probably derived from a fusion of placoid-ganoid scales. The inner part of the scales is made of dense

3147:"Immunodetection of amelogenin-like proteins in the ganoine of experimentally regenerating scales of Calamoichthys calabaricus, a primitive actinopterygian fish" 1028:

along their outer or posterior edges. Because of these teeth, the scales have a rough texture. They are usually found on fishes with spiny fin rays, such as the

2244:) have smooth skin without scales and without dermal bone. Lampreys get some protection from a tough leathery skin. Hagfish exude copious quantities slime or 776:

Cycloid (circular) scales have a smooth texture and are uniform, with a smooth outer edge or margin. They are most common on fish with soft fin rays, such as

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elongate. These penetrate the compact layer of the mesenchyme, which consolidates prior to the formation of the scale, in order to initiate the dermal plate.

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have ctenoid scales on the eyed side and cycloid scales on the blind side, while other species have ctenoid scales in males and cycloid scales in females.

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lack scales and have a layer of smelly and bitter slime which blocks out disease and probably discourages predators, implying their bright coloration is

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form a delimiting layer, which is located in the upper part of the mesenchyme. Subsequently, these cells will differentiate in the area where the scale

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Bone, a tissue that is both resistant to mechanical damage and relatively prone to fossilization, often preserves internal detail, which allows the

1140: 4227:"Modeling how shark and dolphin skin patterns control transitional wall-turbulence vorticity patterns using spatiotemporal phase reset mechanisms" 4164:

Muthuramalingam, Muthukumar; Villemin, Leo S.; Bruecker, Christoph (29 April 2019). "Streak formation in flow over Biomimetic Fish Scale Arrays".

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which have modified cosmoid scales that lack cosmine and are thinner than true cosmoid scales. They are also present in some tetrapodomorphs like

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scales composed of a layer of dense, lamellar collagen bone called isopedine, above which is a layer of tubercles usually composed of bone, as in

4057:"Surface drag reduction and flow separation control in pelagic vertebrates, with implications for interpreting scale morphologies in fossil taxa" 1699:. In laymen's terms, fouling is known as the process by which something becomes encrusted with material from the surrounding environment such as 2080:

that detect water movements. In bony fishes, the scales along the lateral line have central pores that allow water to contact the sensory cells.

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Placoid scales as viewed through an electron microscope. Also called dermal denticles, these are structurally homologous with vertebrate teeth.

1977:, the level of ApoE transcription is high, which has led to the conclusion that this protein is important for the late development of scales. 1715:

have shown that if a biomimetic material can be engineered, it could potentially lead to fuel cost savings for military vessels of up to 45%.

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On the crown of the fast-swimming sharks there are a series of parallel riblets or ridges which run from an anterior to posterior direction.

714: 1897:. Morphogenesis begins at the time when fibroblasts are relocated to the upper part of the compact mesenchyme. Throughout this process, the 3388: 263:

confuse things further, scale morphologies are not unique to taxa, and may be indistinguishable on the same area of two different species.

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Sire, J.Y.; Huysseune, A.N.N. (2003). "Formation of dermal skeletal and dental tissues in fish: a comparative and evolutionary approach".

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Feld, Katrine; Kolborg, Anne Noer; Nyborg, Camilla Marie; Salewski, Mirko; Steffensen, John Fleng; Berg-Sørensen, Kirstine (24 May 2019).

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Turner, S.; Tarling, D. H. (1982). "Thelodont and other agnathan distributions as tests of Lower Paleozoic continental reconstructions".

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are also involved in scale development. The placoid scales of cartilaginous fishes are also called dermal denticles and are structurally

5854: 4959:, gen. et sp. nov., a small and eyeless banjo catfish (Siluriformes: Aspredinidae) from the river channels of the lower Amazon basin". 1473:

Rhomboidal scales with the properties of both placoid and ganoid scales are suspected to exist in modern jawed fish ancestors: jawless

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or slime which can protect against pathogens such as bacteria, fungi, and viruses, and reduce surface resistance when the fish swims.

3163: 3146: 243:, are well understood. The scales were formed and shed throughout the organisms' lifetimes, and quickly separated after their death. 5884: 4891: 4039:"Experimental investigations on drag-reduction characteristics of bionic surface with water-trapping microstructures of fish scales" 3882:

Hage, W.; Bruse, M.; Bechert, D. W. (1 May 2000). "Experiments with three-dimensional riblets as an idealized model of shark skin".

3581: 1748:. One study found that these biomimetic surfaces reduced drag by up to 9%, while with flapping motion drag reduction reached 12.3%. 1363:

is extremely flattened laterally (side to side), leaving the body just millimetres thick, and the body is so silvery as to resemble

5247:"Trophic novelty is linked to exceptional rates of morphological diversification in two adaptive radiations of Cyprinodon pupfish" 2229:

Fish without scales usually evolve alternatives to the protection scales can provide, such as tough leathery skin or bony plates.

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Seahorses lack scales but have thin skin stretched over a bony plate armour arranged in rings through the length of their bodies.

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Domel, August G.; Saadat, Mehdi; Weaver, James C.; Haj-Hariri, Hossein; Bertoldi, Katia; Lauder, George V. (28 February 2018).

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Fish scales with these properties are used in some cosmetics, since they can give a shimmering effect to makeup and lipstick.

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elasmoid scales are used in the lab to study bone mineralization process, and can be cultured (kept) outside of the organism.

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Scales vary enormously in size, shape, structure, and extent, ranging from strong and rigid armour plates in fishes such as

2566: 2518:) is a specialised feeding behaviour in fish that involves eating the scales of other fish. Lepidophagy has independently 341: 5052: 4123:

Lauder, George V.; Wainwright, Dylan K.; Domel, August G.; Weaver, James C.; Wen, Li; Bertoldi, Katia (18 October 2016).

2310:, have developed external bony plates, structurally resembling placoid scales, as protective armour against predators. 5623: 2852:

Turner, S. (1991). "Monophyly and interrelationships of the Thelodonti". In M. M. Chang; Y. H. Liu; G. R. Zhang (eds.).

1536: 458:. They are derived from cosmoid scales and often have serrated edges. They are covered with a layer of hard enamel-like 415: 5523: 5035: 4334:

Liu, Yunhong; Li, Guangji (15 December 2012). "A new method for producing "Lotus Effect" on a biomimetic shark skin".

3034:"Zebrafish as an Emerging Model for Osteoporosis: A Primary Testing Platform for Screening New Osteo-Active Compounds" 2525:

Fish scales can be nutritious, containing a dermal portion and a layer of protein-rich mucus apart from the layers of

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Mesenchyme consolidation: The consolidation or structuring of the mesenchyme originates during the development of the

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for arrow heads, breastplates, and as shielding to cover plows. In current times jewellery is made from these scales.

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Kawasaki, Kenta C., "A Genetic Analysis of Cichlid Scale Morphology" (2016). Masters Theses May 2014 - current. 425.

3514: 1463:-like substance. Placoid scales cannot grow in size, but rather more scales are added as the fish increases in size. 3427: 2325:. Because of this heavy armour boxfish are limited to slow movements, but few other fish are able to eat the adults. 6275: 6002: 5972: 5859: 5468: 1966: 948: 4733:"Epidermal expression of apolipoprotein E gene during fin and scale development and fin regeneration in zebrafish" 2057:

Surgeonfish (left) have a sharp, scalpel-like modified scale on either side just before the tail. Closeup (right).

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Recent research has shown that there is a pre and post-breakdown regime in the near-wall boundary layer where the

1284:. The circumference of the scales grows first, followed by thickness when overlapping layers mineralize together. 6548: 6541: 6508: 6250: 5922: 5603: 3278:

Bruet, B. J. F.; Song, J.; Boyce, M. C.; Ortiz, C. (2008). "Materials design principles of ancient fish armour".

2578: 3597: 1684:, by attaching pieces of shark skin to wooden boards. The small size of the scales grates the food very finely. 1658:-like texture of shark and ray skin, coupled with its toughness, has led it to be valued as a source of rawhide 1229: 6558: 6317: 4587:"Skin development in bony fish with particular emphasis on collagen deposition in the dermis of the zebrafish ( 1822:

have an abdominal row of scutes, which are scales with raised, sharp points that are used for protection. Some

1741: 1330: 1281: 5580: 5084:"Handed Foraging Behavior in Scale-Eating Cichlid Fish: Its Potential Role in Shaping Morphological Asymmetry" 4637:

Sire, J.Y. (2001). "Teeth outside the mouth in teleost fishes: how to benefit from a developmental accident".

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Lauder, George V.; Wainwright, Dylan K.; Domel, August G.; Weaver, James C.; Wen, Li; Bertoldi, Katia (2016).

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are composed of compacted collagen that allow it to maintain the union with the mesenchyme. This are known as

932: 3454: 2795: 6536: 6518: 6080: 5829: 4930: 4685:"Scale development in fish: a review, with description of sonic hedgehog (shh) expression in the zebrafish ( 2906: 2825: 1519:, making swimming both more efficient and quieter compared to that of bony fishes. It also serves a role in 964: 916: 900: 3016:

A microanatomical and histological study of the postcranial dermal skeleton in the Devonian sarcopterygian

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crystallites. Ganoine is an ancient feature of ray-finned fishes, being found for example on the scales of

368:. The layer of dentine that was present in the first lobe-finned fish is usually reduced, as in the extant 6498: 6322: 6307: 6130: 5849: 5738: 2269: 2221: 1950: 1882: 400:, amiids, and teleosts, whose cycloid and ctenoid scales represent the least mineralized elasmoid scales. 382:. Elasmoid scales have appeared several times over the course of fish evolution. They are present in some 121:

of a scale can be used to identify the species of fish it came from. Scales originated within the jawless

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Hori, M. (1993). "Frequency-dependent natural selection in the handedness of scale-eating cichlid fish".

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lack scales, though several families have body armour in the form of dermal plates or some sort of scute.

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and distinctive cellular differentiation makes them exclusive from other structures that arise from the

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Ganoine is a characteristic component of ganoid scales. It is a glassy, often multi-layered mineralized

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and growth of the scales to be studied in detail. The scales comprise a non-growing "crown" composed of

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Diagram of the side profile of a shark denticle showing a vortex in the wake downstream of the denticle

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All denticles are composed of an interior pulp cavity with a nervous and arterial supply rooted in the

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Fletcher, Thomas; Altringham, John; Peakall, Jeffrey; Wignall, Paul; Dorrell, Robert (7 August 2014).

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Dorsal view of right-bending (left) and left-bending (right) jaw morphs adapted for eating fish scales

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Monnot, M.J.; Babin, P.J.; Poleo, G.; Andre, M.; Laforest, L.; Ballagny, C.; Akimenko, M.A. (1999).

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Richter, M. (1995). "A microstructural study of the ganoine tissue of selected lower vertebrates".

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Shono, T.; Thiery, A.P.; Cooper, R.L.; Kurokawa, D.; Britz, R.; Okabe, M.; Fraser, G.J. (2019).

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MICHAEL ALLABY "cosmoid scale ." A Dictionary of Zoology . . Encyclopedia.com. 29 Oct. 2019 <

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Sherman, Vincent R.; Yaraghi, Nicholas A.; Kisailus, David; Meyers, Marc A. (1 December 2016).

2902:"A New Early Devonian thelodont from Celtiberia (Spain), with a revision of Spanish thelodonts" 1725:

Surfaces that mimic the skin of sharks have also been used in order to keep microorganisms and

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of raised ribs (left). The scales themselves are covered with a delicate net pattern (right).

220: 5982: 5927: 5889: 5844: 5802: 5726: 5513: 5447: 5370: 5360: 5313: 5258: 5209: 5113: 5103: 5023: 5006: 4970: 4851: 4843: 4781: 4744: 4700: 4646: 4602: 4542: 4496: 4488: 4423: 4351: 4303: 4262: 4246: 4191: 4144: 4092: 4076: 4004: 3946: 3899: 3828: 3812: 3771: 3753: 3742:"Dermal Denticles of Three Slowly Swimming Shark Species: Microscopy and Flow Visualization" 3709: 3654: 3353: 3337: 3295: 3257: 3215: 3199: 3158: 3100: 3063: 3045: 2974: 2964: 2915: 2882: 2791: 2762: 2707: 2666: 2321:

enclosing the entire body. These bony plates are not modified scales but skin that has been

2039: 1954: 1863: 1755: 1671: 1637: 1467: 1343: 1277: 718: 480: 383: 346: 290: 3500:

Systematics, Zoogeography, and Conservation of the Freshwater Fishes of Peninsular Malaysia

3326:"Microstructural and geometric influences in the protective scales of Atractosteus spatula" 2333: 2048: 6199: 6179: 6007: 5994: 5977: 5932: 5814: 5753: 5677: 5672: 5608: 5595: 5565: 5170: 3387:. Missouri Department of Conservation Fisheries Division. January 22, 2013. Archived from 2603: 2184: 2156: 2106: 2077: 1760: 1642: 1188: 1032:

fishes. These scales contain almost no bone, being composed of a surface layer containing

70: 4650: 4038: 1666:. One of the many historical applications of shark shagreen was in making hand-grips for 1114:

The size of the teeth on ctenoid scales can vary with position, as these scales from the

5309: 5190: 5099: 4839: 4347: 4242: 4187: 4140: 4072: 3895: 3705: 3291: 2960: 2758: 2391:

Some fish, such as hoki and swordfish, are born with scales but shed them as they grow.

1136:

vary from the medial (middle of the fish), to dorsal (top), to caudal (tail end) scales.

544: 6475: 6448: 6340: 6332: 6265: 6235: 6174: 6156: 6110: 6100: 5704: 5657: 5375: 5348: 5118: 5083: 4856: 4823: 4501: 4476: 4267: 4226: 4097: 4056: 3833: 3800: 3776: 3741: 3358: 3325: 3220: 3187: 3068: 3033: 2979: 2944: 2488: 2461: 2340: 2303: 2180: 2152: 2011: 1942: 1935: 1898: 1871: 1540: 1512: 1436: 1364: 1236: 1073: 1033: 750: 641: 492: 396: 378: 364: 302: 4308: 4291: 4008: 3015: 2712: 2695: 2464:, as well as any fish that lack scales (there do not seem to be fish that lack fins). 1105: 6622: 6577: 6480: 6394: 6214: 6189: 6184: 6140: 6135: 6090: 6085: 6065: 5937: 5716: 5635: 5263: 5246: 5010: 4211: 4024: 3919: 3569: 3537: 3203: 2929: 2920: 2901: 2766: 2598: 2538: 2249: 1946: 1851: 1847: 1621: 1204: 1003: 987: 808: 722: 597: 558: 432: 181: 5333: 5280: 5231: 4666: 4562: 3974: 3723: 3128: 2803: 2729: 1917:

occurs, allowing the scale to acquire the rigid characteristic that identifies them.

1881:

Morphogenesis induction: The morphogenesis is due to the formation of the epidermal

1128: 525:

of ray-finned fishes, ganoine or ganoine-like tissues are also found on the extinct

6470: 6294: 6209: 6169: 6070: 6012: 5952: 5879: 5874: 5864: 5780: 5770: 5699: 5682: 5589: 5550: 5543: 5063: 4149: 4124: 3714: 3689: 2886: 2613: 2608: 2530: 2453: 2322: 2233: 2084: 2073: 2002: 1958: 1894: 1827: 1811: 1777: 1745: 1626: 1544: 1524: 1509: 1474: 1460: 1456: 1444: 1440: 1280:

begin to organize themselves in the dermal layer, which leads to the initiation of

1261: 1044:. The enamel of the other scale types is reduced to superficial ridges and ctenii. 971: 738: 534: 522: 473: 326: 314: 208: 126: 45: 1995: 1020:

Ctenoid (toothed) scales are like cycloid scales, except they have small teeth or

359: 5317: 5108: 2969: 1941:

One of the genes that regulate the development of scale formation in fish is the

1367:. The mirrors consist of microscopic structures similar to those used to provide 333:. Cosmoid scales increase in size through the growth of the lamellar bone layer. 6453: 6441: 6367: 6125: 6120: 6095: 6075: 6017: 5912: 5807: 5785: 5775: 5748: 5618: 5570: 4785: 3188:"Origin and evolution of the integumentary skeleton in non-tetrapod vertebrates" 2511: 2318: 2307: 2202:

spines are not modified scales but are developed by an independent gene network.

2102: 1962: 1708: 1600: 1588: 1495: 1322: 1084: 1029: 840: 563: 517: 420: 240: 214: 122: 5452: 5435: 4904: 4847: 4355: 3758: 2259:

are scaleless, though some species are covered with tiny smooth cycloid scales.

1608: 1379:

Most fish in the upper ocean are camouflaged by silvering. In fish such as the

1338: 1300:

scales, where the scale bears spines that are continuous with the scale itself.

876:

Cycloid (circular) scales are usually found on carp-like or salmon-like fishes.

17: 6503: 6270: 6204: 6194: 6115: 6105: 5790: 5743: 5689: 5662: 5645: 4546: 3104: 2671: 2654: 2279: 2273: 2199: 2163: 2142: 1927: 1906: 1902: 1875: 1843: 1823: 1696: 1680: 1584: 1516: 1478: 1433: 1265: 1257: 1253: 1144: 791: 707: 601: 581: 526: 512: 504: 443: 391: 369: 350: 306: 114: 102: 90: 66: 4437: 4363: 4317: 4258: 4088: 4016: 3958: 3911: 3824: 3767: 3666: 3349: 3211: 3112: 3059: 3050: 2352:

seem scaleless, but some species are covered with tiny smooth cycloid scales.

6260: 6245: 6230: 6060: 5721: 5667: 5652: 5630: 5613: 5560: 2637: 2618: 2448: 2117: 2105:

have a scalpel-like blade, which is a modified scale, on either side of the

2088: 1886: 1867: 1839: 1819: 1655: 1395: 1269: 1220: 1080: 862: 768:

Leptoid scales come in two forms: cycloid (smooth) and ctenoid (comb-like).

753:. The scales of some species exhibit bands of uneven seasonal growth called 702: 404: 255: 247: 236: 5384: 5325: 5272: 5223: 5127: 5026:

Northeast Fisheries Science Center, NOAA Fisheries. Accessed 4 August 2019.

4865: 4793: 4758: 4714: 4658: 4616: 4554: 4510: 4492: 4445: 4371: 4276: 4203: 4106: 3966: 3842: 3816: 3785: 3674: 3367: 3341: 3307: 3261: 3229: 3120: 3077: 2988: 2721: 2680: 2252:, scraping off the slime as they go and freeing themselves from a predator. 2224:

lack scales and protect themselves with a layer of smelly and bitter slime.

1550:

Unlike bony fish, sharks have a complicated dermal corset made of flexible

571: 5365: 4705: 4684: 4607: 3903: 3172: 2820:

Janvier, Philippe (1998). "Early vertebrates and their extant relatives".

1729:

from coating the hulls of submarines and ships. One variety is traded as "

1451:, all of which sits on top of a rectangular basal plate that rests on the 1326:

The herring's reflectors are nearly vertical for camouflage from the side.

6463: 6404: 6350: 6345: 6164: 6032: 5694: 5640: 5555: 4750:

10.1002/(SICI)1097-0177(199903)214:3<207::AID-AJA4>3.0.CO;2-5

4412:"The hydrodynamic function of shark skin and two biomimetic applications" 4290:

Magin, Chelsea M.; Cooper, Scott P.; Brennan, Anthony B. (1 April 2010).

3186:

Sire, Jean-Yves; Donoghue, Philip C. J.; Vickaryous, Matthews K. (2009).

2544: 2418: 2402: 2383: 2295: 2291: 2283: 2121: 2096: 1931: 1930:, thus forming bone. The patches of the matrix of the scale that are not 1700: 1663: 1551: 1421: 1311: 1294:

scales, where the margin of the scale bears indentations and projections.

1041: 617: 488: 439: 387: 373: 310: 294: 227: 173: 158: 142: 4080: 3164:

10.1002/(SICI)1097-0185(199709)249:1<86::AID-AR11>3.0.CO;2-X

2282:

have loose, thin skin often covered with fine forked dermal prickles or

1965:, has an interaction with shh, because ApoE provides cholesterol to the 1838:

Scales typically appear late in the development of fish. In the case of

1695:

In the marine industry, there is an extremely large market and need for

1489: 39: 6436: 6431: 6409: 6382: 6377: 6372: 5917: 5869: 5763: 5533: 5528: 4982: 4428: 4411: 4195: 3950: 2552: 2526: 2519: 2368: 2299: 2263: 2241: 2237: 2133: 2113: 2092: 2065: 2005:. Modified scales along the lateral line are visible in the lower half. 1736:

A lot of the new methods for replicating shark skin involve the use of

1659: 1448: 1380: 1372: 1115: 1021: 734: 508: 484: 468: 463: 459: 330: 322: 318: 279: 251: 161:) are covered with placoid scales. Some species are covered instead by 106: 5214: 4824:"Evolution and Developmental Diversity of Skin Spines in Pufferfishes" 4470: 4468: 4250: 4118: 4116: 3735: 3733: 3658: 1248:

Ctenoid scales, similar to other epidermal structures, originate from

6355: 4961: 3986: 3984: 3299: 2780:

Märss, T. (2006). "Exoskeletal ultrasculpture of early vertebrates".

2492: 2480: 2476: 2019: 1890: 1859: 1675: 1531: 1505: 1452: 1273: 939: 891: 812: 777: 733:

Leptoid (bony-ridge) scales are found on higher-order bony fish, the

654: 637: 500: 455: 451: 298: 189: 177: 130: 4974: 1264:

of the fish. The development process begins with an accumulation of

4178: 6458: 6387: 5797: 4477:"Shark skin-inspired designs that improve aerodynamic performance" 4125:"Structure, biomimetics, and fluid dynamics of fish skin surfaces" 3690:"Structure, biomimetics, and fluid dynamics of fish skin surfaces" 2502: 2472: 2371:

have plates of ossified skin fused together to form a rigid shell.

2245: 1815: 1785: 1771: 1726: 1704: 1686: 1667: 1607: 1599: 1558: 1554: 1488: 1417: 1400: 1337: 1329: 1321: 1133: 1127: 1064: 923: 907: 741: 648: 426: 414: 340: 278: 207: 197: 162: 154: 138: 38: 5191:"Functional morphology of feeding in the scale-eating specialist 2696:"Fish scale development: Hair today, teeth and scales yesterday?" 745:

further scales are added in concentric layers as the fish grows.

6360: 6302: 5947: 5500: 5491: 2522:

in at least five freshwater families and seven marine families.

2434: 2398: 2146: 1539:

of the shark. Both riblet shapes assist in creating a turbulent

1172: 1155:

Ctenoid (toothed) scales are usually found on perch-like fishes.

781: 313:

bone called isopedine. On top of this lies a layer of spongy or

196:

with vertebrate teeth. Most fish are also covered in a layer of

185: 165:, and others have no outer covering on part or all of the skin. 134: 62: 49: 5464: 3032:

Bergen, Dylan J. M.; Kague, Erika; Hammond, Chrissy L. (2019).

3002: 2018:

Closeup of a modified cycloid scale from the lateral line of a

1850:

is induced, and finally the process of differentiation or late

865:

fish with circular cycloid scales that are smooth to the touch.

93: 6399: 5967: 4225:

Bandyopadhyay, Promode R.; Hellum, Aren M. (23 October 2014).

3991:

Dou, Zhaoliang; Wang, Jiadao; Chen, Darong (1 December 2012).

2854:

Early Vertebrates and Related Problems of Evolutionary Biology

2349: 2317:

In boxfish, the plates fuse together to form a rigid shell or

2256: 1306:

scales, where the spines on the scale are distinct structures.

1147:

have cycloid scales on the belly but ctenoid scales elsewhere.

496: 447: 146: 110: 1945:(shh) gene, which by means of the (shh) protein, involved in 1803:

a modified, thickened scale that often is keeled or spiny, or

1334:

The deep sea hatchetfish has scales which reflect blue light.

1087:

fish with toothed ctenoid scales that are rough to the touch.

3502:(Doctoral Dissertation ed.). Colorado State University. 2386:

have thin skin stretched over bony plates arranged in rings.

1310:

Most ray-finned fishes have ctenoid scales. Some species of

5436:"Scales, enamel, cosmine, ganoine, and early osteichthyans" 1439:("denticle" translates to "small tooth"), having a central 1287:

Ctenoid scales can be further subdivided into three types:

3382:"Missouri Alligator Gar Management and Restoration Plan" 2867:

Märss, T. (1986). "Squamation of the thelodont agnathan

2796:

10.1671/0272-4634(2006)26[235:EUOEV]2.0.CO;2

2068:

a number of modified scales to serve various functions.

676:

Earrings made from the ganoid scales of an alligator gar

317:

bone supplied with blood vessels, followed by a complex

85:, as well as possible hydrodynamic advantages. The term 5460: 3805:

Proceedings of the Royal Society B: Biological Sciences

3551:

The Diversity of Fishes Biology, Evolution, and Ecology

1466:

Similar scales can also be found under the head of the

4585:

Le Guellec, D.; Morvan-Dubois, G.; Sire, J.Y. (2004).

3412:

Lagler, K. F., J. E. Bardach, and R. R. Miller (1962)

2248:

if they are threatened. They can tie themselves in an

2128:, which are easily shed and aid in escaping predators. 1926:

cavities. At this point, elasmoblasts are replaced by

431:

Mineral texture of ganoine layers in the scales of an

4410:

Lauder, George V.; Oeffner, Johannes (1 March 2012).

3576:. Oxford: Oxford University Press. pp. 193–195. 2900:

Botella, H.; J. I. Valenzuela-Rios; P. Carls (2006).

1470:. The amount of scale coverage is much less in rays. 5401:

Helfman, G.S., B.B. Collette and D.E. Facey (1997).

4329: 4327: 3429:

Exotic Animal Medicine for the Veterinary Technician

1957:(ApoE), that allows the transport and metabolism of 811:

have large cycloid scales arranged on the fish in a

537:

in vertebrates or even considered a type of enamel.

6491: 6424: 6331: 6293: 6284: 6223: 6154: 6041: 5993: 5903: 5828: 5588: 5498: 4882:. In: McKenzie DJ, Farrell AP and Brauner CJ (Eds) 4457: 4455: 3993:"Bionic Research on Fish Scales for Drag Reduction" 2421:

appear scaleless because their scales are so small.

3931: 3929: 3538:http://scholarworks.umass.edu/masters_theses_2/425 3014:Zylberberg, L., Meunier, F.J., Laurin, M. (2010). 4386:"Sharklet Discovery | Sharklet Technologies, Inc" 4055:Palmer, Colin; Young, Mark T. (14 January 2015). 3598:"There Are Probably Fish Scales In Your Lipstick" 3140: 3138: 2746:Palaeogeography, Palaeoclimatology, Palaeoecology 1814:, are completely or partially covered in scutes. 3145:Zylberberg, L.; Sire, J. -Y.; Nanci, A. (1997). 1670:. The rough texture of the skin is also used in 1412:(pointed, tooth-shaped) scales are found in the 1342:The scales of a typical teleost fish, like this 271:extensive comparison between ecological niches. 5184: 5182: 5180: 3877: 3875: 1969:. It has been shown that during the process of 688:Fossil of a primitive rayfin with ganoid scales 4693:International Journal of Developmental Biology 4595:International Journal of Developmental Biology 4580: 4578: 4576: 4574: 4572: 3564: 3562: 3560: 3319: 3317: 3243: 3241: 3239: 2560:. The jaws of this fish occur in two distinct 2437:seem largely free of scales, but they are not. 5476: 5047: 5045: 5043: 4726: 4724: 3516:The Fishes of the Indo-Australian Archipelago 2943:Ferrón, Humberto G.; Botella, Héctor (2017). 2815: 2813: 109:, to microscopic or absent in fishes such as 61:is a small rigid plate that grows out of the 8: 5425:: CS1 maint: multiple names: authors list ( 4905:"Hagfish slime: The clothing of the future?" 4878:Coolidge E, Hedrick MS and Milsom WK (2011) 4678: 4676: 3426:Ballard, Bonnie; Cheek, Ryan (2 July 2016). 3273: 3271: 2847: 2845: 2551:. An exception occurs with the scale-eating 466:, and a layer of inorganic bone salt called 125:, ancestors to all jawed fishes today. Most 4632: 4630: 4628: 4626: 4528: 4526: 4524: 4522: 4520: 2087:and chimaeras, the stinging tail spines of 1604:Effects of turbulent flow on boundary layer 1067:has a toothed outer edge (at top of image). 6290: 5483: 5469: 5461: 5082:Lee, H. J.; Kusche, H.; Meyer, A. (2012). 5077: 5075: 2856:. Science Press, Beijing. pp. 87–119. 2653:Mongera, A.; Nüsslein-Volhard, C. (2013). 1953:, enable the formation of the scales. The 1870:substrate organized by perpendicularly by 1620:(COT) through multiple different avenues. 843:has a smooth outer edge (at top of image). 653:Geometrically laid out ganoid scales on a 188:involved in tooth and hair development in 6028:Tradeoffs for locomotion in air and water 5451: 5374: 5364: 5347:Stewart, T. A.; Albertson, R. C. (2010). 5262: 5213: 5117: 5107: 4855: 4748: 4704: 4606: 4500: 4427: 4307: 4266: 4177: 4148: 4096: 3832: 3775: 3757: 3713: 3357: 3250:Zoological Journal of the Linnean Society 3219: 3162: 3067: 3049: 2978: 2968: 2919: 2711: 2670: 2155:have scales modified into large external 1789:superficially similar to that of scales. 1498:, have placoid scales (dermal denticles). 1371:: stacks of between 5 and 10 crystals of 289:Cosmoid scales are found only on ancient 27:Rigid covering growing atop a fish's skin 4336:Journal of Colloid and Interface Science 1150: 1139: 871: 620:has rows of ganoid scales enlarged into 539: 4807:How the pufferfish got its wacky spines 4774:Molecular Phylogenetics & Evolution 3858:"The Importance of Being Cartilaginous" 2630: 2407: 2401:only along the lateral line and in the 2358: 2211: 2170: 1161: 1112: 880: 805: 666: 586: 483:that covers the scales, as well as the 129:are covered with the cycloid scales of 5418: 5166: 5156: 4931:"No One Is Prepared for Hagfish Slime" 4481:Journal of the Royal Society Interface 3330:Journal of the Royal Society Interface 2945:"Squamation and ecology of thelodonts" 2290:Many groups of bony fishes, including 2099:are fused and modified placoid scales. 1800:an external shield-like bony plate, or 1587:. During a recent research experiment 1040:and a deeper layer composed mostly of 386:, such as all extant and some extinct 6608:Hydrodynamic aspects of shark scales 5405:. Blackwell Science. pp. 33–36. 3860:. ReefQuest Centre for Shark Research 3532: 3530: 3528: 3526: 2347: 1455:. The outermost layer is composed of 7: 5245:Martin, C.; P.C. Wainwright (2011). 4651:10.1046/j.1525-142x.2001.003002104.x 3453:McGrouther, Mark (2 December 2019). 2655:"Scales of fish arise from mesoderm" 2495:, snake mackerels, and puffer fish. 1885:, which is generated by joining the 721:used the tough ganoid scales of the 325:with a superficial outer coating of 6584: 5855:Electroreception and electrogenesis 5202:The Journal of Experimental Biology 4955:Friel, J P; Lundberg, J G (1996). " 4683:Sire, J.Y.; Akimenko, M.A. (2003). 4166:The Journal of Experimental Biology 1826:have a row of scutes following the 1547:farther away from the sharks skin. 1447:, surrounded by a conical layer of 661: 529:. It has been suggested ganoine is 372:, or entirely absent, as in extant 168:Fish scales are part of the fish's 73:, which can also provide effective 6613:Fish scales and flow manipulation 5141:Froese, R. and D. Pauly. Editors. 4292:"Non-toxic antifouling strategies" 3416:. New York: John Wiley & Sons. 2874:Journal of Vertebrate Paleontology 2783:Journal of Vertebrate Paleontology 1922: 1874:. Subsequently, for both fish the 1428:. Placoid scales are structurally 25: 4903:Rothschild, Anna (1 April 2013). 4884:Fish Physiology: Primitive Fishes 1616:Sharks decrease drag and overall 640:are reduced in size and resemble 293:, including some of the earliest 69:is covered with these protective 6583: 6572: 6571: 6554: 6553: 5579: 5264:10.1111/j.1558-5646.2011.01294.x 5011:10.1111/j.0022-1112.2005.00692.x 3801:"Hydrodynamics of fossil fishes" 3204:10.1111/j.1469-7580.2009.01046.x 2921:10.1111/j.1475-4983.2005.00534.x 2426: 2410: 2376: 2361: 2339: 2332: 2214: 2192: 2173: 2047: 2038: 2010: 1994: 1494:Cartilaginous fishes, like this 1228: 1212: 1196: 1180: 1164: 1104: 1097: 1072: 1056: 995: 979: 963: 947: 931: 915: 899: 883: 848: 832: 797: 790: 693: 681: 669: 629: 609: 589: 570: 543: 180:, which distinguishes them from 5539:Environmental impact of fishing 4416:Journal of Experimental Biology 3519:. E.J. Brill. pp. 306–307. 3020:, Acta Palaeontologica Polonica 2137:darters have a row of enlarged 438:Ganoid scales are found in the 98:, meaning a shell pod or husk. 4150:10.1103/PhysRevFluids.1.060502 3715:10.1103/PhysRevFluids.1.060502 2887:10.1080/02724634.1986.10011593 2064:Different groups of fish have 1256:. Development starts near the 761:). These bands can be used to 700:Ganoid scales on a fossilised 254:, with a sometimes-ornamented 1: 4309:10.1016/S1369-7021(10)70058-4 4009:10.1016/S1672-6529(11)60140-6 3997:Journal of Bionic Engineering 3574:The Biology of the Deep Ocean 2713:10.1016/S0960-9822(01)00438-9 2567:frequency-dependent selection 2397:Prominent scaling appears on 557:Ganoid scales of the extinct 507:. It is composed of rod-like 423:appear glassy due to ganoine. 5624:intramembranous ossification 5318:10.1126/science.260.5105.216 5109:10.1371/journal.pone.0044670 4929:Yong, Ed (23 January 2019). 3455:"Cycloid and Ctenoid Scales" 3003:https://www.encyclopedia.com 2970:10.1371/journal.pone.0172781 2767:10.1016/0031-0182(82)90027-X 2539:much smaller than their prey 2533:. They are a rich source of 2479:, for a fish to be declared 1796:, and can take the form of: 1691:Barnacle growth on boat hull 239:, the most abundant form of 212:Left to right: denticles of 172:, and are produced from the 65:of a fish. The skin of most 4786:10.1016/j.ympev.2013.03.014 4639:Evolution & Development 2485:kosher certification agency 2338: 1792:Scute comes from Latin for 1744:properties, exhibiting the 1718:There are many examples of 1153: 1103: 874: 796: 621: 556: 521:. While often considered a 137:, or the ctenoid scales of 44:Cycloid scales cover these 32:Fish scale (disambiguation) 6645: 6003:Fin and flipper locomotion 5973:Sequential hermaphroditism 5860:Jamming avoidance response 5577: 5453:10.1016/j.crpv.2015.04.001 5069:. Retrieved 9 August 2019. 4886:, Elsevier, Page 182–213. 4848:10.1016/j.isci.2019.06.003 4356:10.1016/j.jcis.2012.08.033 4061:Royal Society Open Science 3759:10.3390/biomimetics4020038 3093:Osteoporosis International 3038:Frontiers in Endocrinology 2643:. Retrieved 28 April 2019. 2306:, and several families of 2183:have scales modified into 1393: 358:Elasmoid scales are thin, 141:, or the ganoid scales of 29: 6567: 4547:10.1017/S1464793102006073 3432:. John Wiley & Sons. 3105:10.1007/s00198-013-2441-3 2672:10.1016/j.cub.2013.02.056 2579:Age determination in fish 2471:or divine decrees of the 2091:, and the "saw" teeth of 2083:The dorsal fin spines of 2072:Almost all fishes have a 826:Cycloid (circular) scales 600:has a tough armouring of 3051:10.3389/fendo.2019.00006 1921:Unlike elasmoid scales, 1866:. Bony fish generate an 1394:Not to be confused with 1050:Ctenoid (toothed) scales 6519:Glossary of ichthyology 6081:Diel vertical migration 5434:Schultze, H.P. (2016). 5403:The Diversity of Fishes 5189:Janovetz, Jeff (2005). 5143:"Glossary: Lepidophagy" 5067:OU Kosher Certification 5053:"All About Kosher Fish" 4999:Journal of Fish Biology 4462:10.1098/rsta.2010.0201. 2826:Oxford University Press 1424:. They are also called 1063:The ctenoid scale of a 839:The cycloid scale of a 719:people of the Caribbean 636:The ganoid scales on a 580:Most ganoid scales are 523:synapomorphic character 353:, have elasmoid scales. 329:. The upper surface is 305:, including the living 94: 5885:Surface wave detection 5850:Hydrodynamic reception 5524:Diseases and parasites 5440:Comptes Rendus Palevol 4737:Developmental Dynamics 4493:10.1098/rsif.2017.0828 4129:Physical Review Fluids 3817:10.1098/rspb.2014.0703 3694:Physical Review Fluids 3549:Helfman, Gene (2009). 3342:10.1098/rsif.2016.0595 3262:10.1006/zjls.1995.0023 2694:Sharpe, P. T. (2001). 2621:– Japanese fish scaler 2508: 2433:To casual examination 1951:cellular communication 1949:and in the process of 1782: 1780:are covered in scutes. 1692: 1613: 1605: 1500: 1459:, a largely inorganic 1406: 1347: 1335: 1327: 1148: 1137: 1132:Ctenoid scales from a 657: 604:-shaped ganoid scales. 435: 424: 355: 349:, like this preserved 286: 232: 54: 6023:Undulatory locomotion 5840:Ampullae of Lorenzini 5366:10.1186/1741-7007-8-8 5064:Verifying Kosher Fish 5024:Do tunas have scales? 4880:"Ventilatory Systems" 4706:10.1387/ijdb.15272389 4608:10.1387/ijdb.15272388 3939:Journal of Morphology 3904:10.1007/s003480050400 3884:Experiments in Fluids 3459:The Australian Museum 3151:The Anatomical Record 3018:Eusthenopteron foordi 2557:Perissodus microlepis 2506: 1967:shh signaling pathway 1915:matrix mineralization 1893:through a process of 1775: 1697:anti-fouling surfaces 1690: 1650:Technical application 1611: 1603: 1492: 1404: 1369:structural coloration 1341: 1333: 1325: 1143: 1131: 858:Poropuntius huguenini 652: 430: 418: 344: 282: 211: 42: 6251:Genetically modified 5208:(Pt 24): 4757–4768. 4172:(Pt 16): jeb205963. 2828:. pp. 123–127. 2001:Cycloid scales of a 1971:cell differentiation 1738:polydimethylsiloxane 1720:biomimetic materials 1414:cartilaginous fishes 1119:Cetonurus crassiceps 564:Amblypterus striatus 552:Amblypterus striatus 491:in some non-teleost 376:and in the Devonian 170:integumentary system 151:Cartilaginous fishes 30:For other uses, see 6056:Aquatic respiration 5943:Life history theory 5310:1993Sci...260..216H 5100:2012PLoSO...744670L 4840:2019iSci...19.1248S 4348:2012JCIS..388..235L 4243:2014NatSR...4E6650B 4188:2019arXiv190412752M 4141:2016PhRvF...1f0502L 4081:10.1098/rsos.140163 4073:2015RSOS....240163P 3896:2000ExFl...28..403B 3706:2016PhRvF...1f0502L 3625:"Skin of the Teeth" 3513:E.J. Brill (1953). 3498:Ismail, M. (1989). 3292:2008NatMa...7..748B 2961:2017PLoSO..1272781F 2759:1982PPP....39..295T 2514:(Ancient Greek for 2208:Fish without scales 1810:Some fish, such as 1756:Parametric modeling 419:The scales of this 321:-like layer called 284:Queensland lungfish 235:The bony scales of 77:through the use of 5895:Weberian apparatus 5169:has generic name ( 4535:Biological Reviews 4429:10.1242/jeb.063040 4231:Scientific Reports 4196:10.1242/jeb.205963 3951:10.1002/jmor.20047 3856:Martin, R. Aidan. 3811:(1788): 20140703. 3623:Martin, R. Aidan. 3553:. Wiley-Blackwell. 3192:Journal of Anatomy 2549:bilateral symmetry 2509: 1783: 1693: 1614: 1606: 1523:by exhibiting the 1501: 1407: 1361:marine hatchetfish 1348: 1336: 1328: 1149: 1138: 658: 436: 425: 384:lobe-finned fishes 356: 347:Lobe-finned fishes 291:lobe-finned fishes 287: 233: 55: 6599: 6598: 6509:Fish common names 6420: 6419: 6051:Aquatic predation 5875:Capacity for pain 5604:Age determination 5412:978-0-86542-256-8 5304:(5105): 216–219. 5215:10.1242/jeb.01938 5036:Leviticus 11:9–10 4957:Micromyzon akamai 4487:(139): 20170828. 4251:10.1038/srep06650 3945:(10): 1096–1110. 3659:10.1021/la0401011 3439:978-1-118-92421-1 3336:(125): 20160595. 2835:978-0-19-854047-2 2822:Early Vertebrates 2706:(18): R751–R752. 2594:Photonic crystals 2589:Animal reflectors 2584:Animal coloration 2535:calcium phosphate 2467:According to the 2357: 2356: 1864:mesenchymal cells 1834:Scale development 1618:cost of transport 1246: 1245: 1126: 1125: 1038:calcium carbonate 1013: 1012: 821: 820: 816: 578: 577: 515:actinopteryigian 493:ray-finned fishes 390:, as well as the 89:derives from the 16:(Redirected from 6636: 6587: 6586: 6575: 6574: 6557: 6556: 6291: 5583: 5514:Ethnoichthyology 5485: 5478: 5471: 5462: 5457: 5455: 5430: 5424: 5416: 5389: 5388: 5378: 5368: 5344: 5338: 5337: 5291: 5285: 5284: 5266: 5257:(8): 2197–2212. 5242: 5236: 5235: 5217: 5199: 5186: 5175: 5174: 5168: 5164: 5162: 5154: 5152: 5150: 5138: 5132: 5131: 5121: 5111: 5079: 5070: 5061: 5055: 5049: 5038: 5033: 5027: 5021: 5015: 5014: 5005:(5): 1411–1418. 4993: 4987: 4986: 4952: 4946: 4945: 4943: 4941: 4926: 4920: 4919: 4917: 4915: 4900: 4894: 4876: 4870: 4869: 4859: 4819: 4813: 4804: 4798: 4797: 4769: 4763: 4762: 4752: 4728: 4719: 4718: 4708: 4699:(2–3): 233–247. 4680: 4671: 4670: 4634: 4621: 4620: 4610: 4601:(2–3): 217–231. 4582: 4567: 4566: 4530: 4515: 4514: 4504: 4472: 4463: 4459: 4450: 4449: 4431: 4407: 4401: 4400: 4398: 4396: 4390:www.sharklet.com 4382: 4376: 4375: 4331: 4322: 4321: 4311: 4287: 4281: 4280: 4270: 4222: 4216: 4215: 4181: 4161: 4155: 4154: 4152: 4120: 4111: 4110: 4100: 4052: 4046: 4045: 4043: 4035: 4029: 4028: 3988: 3979: 3978: 3933: 3924: 3923: 3879: 3870: 3869: 3867: 3865: 3853: 3847: 3846: 3836: 3796: 3790: 3789: 3779: 3761: 3737: 3728: 3727: 3717: 3685: 3679: 3678: 3642: 3636: 3635: 3633: 3631: 3620: 3614: 3613: 3611: 3609: 3594: 3588: 3587: 3566: 3555: 3554: 3546: 3540: 3534: 3521: 3520: 3510: 3504: 3503: 3495: 3489: 3488: 3476: 3470: 3469: 3467: 3465: 3450: 3444: 3443: 3423: 3417: 3410: 3404: 3403: 3401: 3399: 3393: 3386: 3378: 3372: 3371: 3361: 3321: 3312: 3311: 3300:10.1038/nmat2231 3280:Nature Materials 3275: 3266: 3265: 3245: 3234: 3233: 3223: 3183: 3177: 3176: 3166: 3142: 3133: 3132: 3088: 3082: 3081: 3071: 3053: 3029: 3023: 3012: 3006: 2999: 2993: 2992: 2982: 2972: 2940: 2934: 2933: 2923: 2897: 2891: 2890: 2864: 2858: 2857: 2849: 2840: 2839: 2817: 2808: 2807: 2777: 2771: 2770: 2753:(3–4): 295–311. 2740: 2734: 2733: 2715: 2691: 2685: 2684: 2674: 2665:(9): R338–R339. 2650: 2644: 2635: 2430: 2414: 2380: 2365: 2343: 2336: 2329: 2328: 2218: 2196: 2177: 2153:Porcupine fishes 2126:deciduous scales 2078:mechanoreceptors 2051: 2042: 2014: 1998: 1955:apolipoprotein E 1830:on either side. 1761:angles of attack 1742:superhydrophobic 1672:Japanese cuisine 1566:Scale morphology 1468:denticle herring 1426:dermal denticles 1344:Atlantic herring 1278:Collagen fibrils 1232: 1216: 1200: 1184: 1168: 1151: 1108: 1101: 1094: 1093: 1076: 1060: 999: 983: 967: 951: 935: 919: 903: 887: 872: 852: 836: 807: 801: 794: 787: 786: 715:Native Americans 697: 685: 673: 633: 613: 593: 574: 547: 540: 462:in the place of 204:Thelodont scales 97: 21: 6644: 6643: 6639: 6638: 6637: 6635: 6634: 6633: 6619: 6618: 6605: 6600: 6595: 6563: 6487: 6416: 6327: 6280: 6219: 6150: 6043: 6037: 5989: 5933:Ichthyoplankton 5899: 5831: 5824: 5820:Digital Library 5815:Teleost leptins 5754:Shark cartilage 5678:pharyngeal slit 5673:pharyngeal arch 5609:Anguilliformity 5594: 5592: 5584: 5575: 5494: 5489: 5446:(1–2): 83–102. 5433: 5417: 5413: 5400: 5397: 5395:Further reading 5392: 5346: 5345: 5341: 5293: 5292: 5288: 5244: 5243: 5239: 5197: 5193:Catoprion mento 5188: 5187: 5178: 5165: 5155: 5148: 5146: 5140: 5139: 5135: 5081: 5080: 5073: 5062: 5058: 5050: 5041: 5034: 5030: 5022: 5018: 4995: 4994: 4990: 4975:10.2307/1447528 4954: 4953: 4949: 4939: 4937: 4928: 4927: 4923: 4913: 4911: 4902: 4901: 4897: 4877: 4873: 4821: 4820: 4816: 4812:, 25 July 2019. 4805: 4801: 4771: 4770: 4766: 4730: 4729: 4722: 4682: 4681: 4674: 4636: 4635: 4624: 4584: 4583: 4570: 4532: 4531: 4518: 4474: 4473: 4466: 4460: 4453: 4409: 4408: 4404: 4394: 4392: 4384: 4383: 4379: 4333: 4332: 4325: 4296:Materials Today 4289: 4288: 4284: 4224: 4223: 4219: 4163: 4162: 4158: 4122: 4121: 4114: 4054: 4053: 4049: 4041: 4037: 4036: 4032: 3990: 3989: 3982: 3935: 3934: 3927: 3881: 3880: 3873: 3863: 3861: 3855: 3854: 3850: 3798: 3797: 3793: 3739: 3738: 3731: 3687: 3686: 3682: 3644: 3643: 3639: 3629: 3627: 3622: 3621: 3617: 3607: 3605: 3604:. 23 April 2015 3596: 3595: 3591: 3584: 3568: 3567: 3558: 3548: 3547: 3543: 3535: 3524: 3512: 3511: 3507: 3497: 3496: 3492: 3478: 3477: 3473: 3463: 3461: 3452: 3451: 3447: 3440: 3425: 3424: 3420: 3411: 3407: 3397: 3395: 3391: 3384: 3380: 3379: 3375: 3323: 3322: 3315: 3277: 3276: 3269: 3247: 3246: 3237: 3185: 3184: 3180: 3144: 3143: 3136: 3090: 3089: 3085: 3031: 3030: 3026: 3013: 3009: 3000: 2996: 2955:(2): e0172781. 2942: 2941: 2937: 2899: 2898: 2894: 2866: 2865: 2861: 2851: 2850: 2843: 2836: 2819: 2818: 2811: 2779: 2778: 2774: 2742: 2741: 2737: 2700:Current Biology 2693: 2692: 2688: 2659:Current Biology 2652: 2651: 2647: 2636: 2632: 2628: 2604:Scale (zoology) 2575: 2501: 2445: 2438: 2431: 2422: 2415: 2387: 2381: 2372: 2366: 2353: 2225: 2219: 2210: 2203: 2197: 2188: 2178: 2139:caducous scales 2107:caudal peduncle 2062: 2061: 2060: 2059: 2058: 2054: 2053: 2052: 2044: 2043: 2034: 2033: 2027: 2026: 2025: 2024: 2023: 2022: 2015: 2007: 2006: 1999: 1990: 1989: 1983: 1981:Modified scales 1872:collagen fibers 1836: 1781: 1770: 1652: 1643:Strouhal arrays 1598: 1568: 1499: 1487: 1477:and then jawed 1399: 1392: 1346:, are silvered. 1320: 1240: 1239: 1233: 1224: 1223: 1217: 1208: 1207: 1201: 1192: 1191: 1185: 1176: 1175: 1169: 1158: 1156: 1122: 1092: 1091: 1090: 1089: 1088: 1077: 1069: 1068: 1061: 1052: 1051: 1018: 1007: 1006: 1000: 991: 990: 984: 975: 974: 968: 959: 958: 952: 943: 942: 936: 927: 926: 920: 911: 910: 904: 895: 894: 888: 877: 870: 869: 868: 867: 866: 853: 845: 844: 837: 828: 827: 817: 774: 731: 710: 698: 689: 686: 677: 674: 645: 634: 625: 614: 605: 594: 554: 549: 548: 413: 354: 339: 337:Elasmoid scales 285: 277: 206: 53: 35: 28: 23: 22: 18:Dermal denticle 15: 12: 11: 5: 6642: 6640: 6632: 6631: 6621: 6620: 6617: 6616: 6611: 6604: 6603:External links 6601: 6597: 6596: 6594: 6593: 6581: 6568: 6565: 6564: 6562: 6561: 6551: 6546: 6545: 6544: 6539: 6531: 6526: 6521: 6516: 6511: 6506: 6501: 6495: 6493: 6489: 6488: 6486: 6485: 6484: 6483: 6478: 6468: 6467: 6466: 6461: 6456: 6446: 6445: 6444: 6439: 6428: 6426: 6422: 6421: 6418: 6417: 6415: 6414: 6413: 6412: 6407: 6402: 6392: 6391: 6390: 6385: 6380: 6375: 6365: 6364: 6363: 6358: 6353: 6348: 6337: 6335: 6333:Wild fisheries 6329: 6328: 6326: 6325: 6320: 6315: 6310: 6305: 6299: 6297: 6288: 6282: 6281: 6279: 6278: 6273: 6268: 6263: 6258: 6256:Hallucinogenic 6253: 6248: 6243: 6238: 6233: 6227: 6225: 6221: 6220: 6218: 6217: 6212: 6207: 6202: 6197: 6192: 6187: 6182: 6177: 6172: 6167: 6161: 6159: 6152: 6151: 6149: 6148: 6143: 6138: 6133: 6131:Schooling fish 6128: 6123: 6118: 6113: 6108: 6103: 6098: 6093: 6091:Filter feeders 6088: 6083: 6078: 6073: 6068: 6066:Bottom feeders 6063: 6058: 6053: 6047: 6045: 6039: 6038: 6036: 6035: 6030: 6025: 6020: 6015: 6010: 6005: 5999: 5997: 5991: 5990: 5988: 5987: 5986: 5985: 5975: 5970: 5965: 5960: 5955: 5950: 5945: 5940: 5935: 5930: 5925: 5920: 5915: 5909: 5907: 5901: 5900: 5898: 5897: 5892: 5887: 5882: 5877: 5872: 5867: 5862: 5857: 5852: 5847: 5842: 5836: 5834: 5826: 5825: 5823: 5822: 5817: 5812: 5811: 5810: 5805: 5795: 5794: 5793: 5788: 5778: 5773: 5768: 5767: 5766: 5756: 5751: 5746: 5741: 5736: 5735: 5734: 5724: 5719: 5714: 5712:Leydig's organ 5709: 5708: 5707: 5705:pharyngeal jaw 5702: 5692: 5687: 5686: 5685: 5680: 5675: 5670: 5665: 5660: 5658:branchial arch 5650: 5649: 5648: 5638: 5633: 5628: 5627: 5626: 5621: 5611: 5606: 5600: 5598: 5586: 5585: 5578: 5576: 5574: 5573: 5568: 5563: 5558: 5553: 5548: 5547: 5546: 5541: 5536: 5526: 5521: 5516: 5511: 5505: 5503: 5496: 5495: 5490: 5488: 5487: 5480: 5473: 5465: 5459: 5458: 5431: 5411: 5396: 5393: 5391: 5390: 5339: 5286: 5237: 5176: 5133: 5071: 5056: 5051:Aryeh Citron, 5039: 5028: 5016: 4988: 4969:(3): 641–648. 4947: 4921: 4895: 4871: 4814: 4799: 4780:(1): 150–160. 4764: 4743:(3): 207–215. 4720: 4672: 4645:(2): 104–108. 4622: 4568: 4541:(2): 219–249. 4516: 4464: 4451: 4422:(5): 785–795. 4402: 4377: 4342:(1): 235–242. 4323: 4282: 4217: 4156: 4112: 4047: 4030: 4003:(4): 457–464. 3980: 3925: 3890:(5): 403–412. 3871: 3848: 3791: 3729: 3680: 3653:(3): 956–961. 3637: 3615: 3602:HuffPost India 3589: 3582: 3570:Herring, Peter 3556: 3541: 3522: 3505: 3490: 3471: 3445: 3438: 3418: 3405: 3394:on May 6, 2016 3373: 3313: 3286:(9): 748–756. 3267: 3256:(2): 173–212. 3235: 3198:(4): 409–440. 3178: 3134: 3099:(2): 567–578. 3083: 3024: 3007: 2994: 2935: 2914:(1): 141–154. 2892: 2859: 2841: 2834: 2809: 2790:(2): 235–252. 2772: 2735: 2686: 2645: 2629: 2627: 2624: 2623: 2622: 2616: 2611: 2606: 2601: 2596: 2591: 2586: 2581: 2574: 2571: 2547:normally show 2500: 2497: 2493:freshwater cod 2489:Orthodox Union 2444: 2441: 2440: 2439: 2432: 2425: 2423: 2416: 2409: 2389: 2388: 2382: 2375: 2373: 2367: 2360: 2355: 2354: 2348: 2345: 2344: 2337: 2327: 2326: 2315: 2288: 2287: 2277: 2267: 2260: 2253: 2227: 2226: 2220: 2213: 2209: 2206: 2205: 2204: 2198: 2191: 2189: 2181:Porcupine fish 2179: 2172: 2169: 2168: 2160: 2150: 2129: 2110: 2100: 2085:dogfish sharks 2081: 2076:, a system of 2056: 2055: 2046: 2045: 2037: 2036: 2035: 2031: 2030: 2029: 2028: 2017: 2016: 2009: 2008: 2000: 1993: 1992: 1991: 1987: 1986: 1985: 1984: 1982: 1979: 1943:sonic hedgehog 1936:Sharpey fibers 1919: 1918: 1910: 1879: 1835: 1832: 1816:River herrings 1808: 1807: 1804: 1801: 1776: 1769: 1766: 1651: 1648: 1597: 1596:Drag reduction 1594: 1585:turbulent flow 1573:stratum laxum. 1567: 1564: 1557:arranged as a 1541:boundary layer 1537:posterior side 1493: 1486: 1483: 1443:supplied with 1391: 1390:Placoid scales 1388: 1365:aluminium foil 1319: 1316: 1308: 1307: 1301: 1295: 1282:mineralization 1244: 1243: 1242: 1241: 1235: 1234: 1227: 1225: 1219: 1218: 1211: 1209: 1203: 1202: 1195: 1193: 1187: 1186: 1179: 1177: 1171: 1170: 1163: 1159: 1154: 1124: 1123: 1113: 1110: 1109: 1102: 1078: 1071: 1070: 1062: 1055: 1054: 1053: 1049: 1048: 1047: 1046: 1034:hydroxyapatite 1017: 1016:Ctenoid scales 1014: 1011: 1010: 1009: 1008: 1002: 1001: 994: 992: 986: 985: 978: 976: 970: 969: 962: 960: 954: 953: 946: 944: 938: 937: 930: 928: 922: 921: 914: 912: 906: 905: 898: 896: 890: 889: 882: 878: 875: 854: 847: 846: 838: 831: 830: 829: 825: 824: 823: 822: 819: 818: 806: 803: 802: 795: 773: 772:Cycloid scales 770: 730: 729:Leptoid scales 727: 712: 711: 699: 692: 690: 687: 680: 678: 675: 668: 662:cycloid scales 647: 646: 642:cycloid scales 635: 628: 626: 624:armour plates. 615: 608: 606: 595: 588: 576: 575: 568: 555: 550: 412: 409: 397:Eusthenopteron 379:Eusthenopteron 365:Eusthenopteron 345: 338: 335: 303:Crossopterygii 283: 276: 275:Cosmoid scales 273: 205: 202: 182:reptile scales 43: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 6641: 6630: 6627: 6626: 6624: 6615: 6612: 6610: 6607: 6606: 6602: 6592: 6591: 6582: 6580: 6579: 6570: 6569: 6566: 6560: 6559:more lists... 6552: 6550: 6547: 6543: 6540: 6538: 6535: 6534: 6532: 6530: 6527: 6525: 6522: 6520: 6517: 6515: 6514:Fish families 6512: 6510: 6507: 6505: 6502: 6500: 6499:Aquarium life 6497: 6496: 6494: 6490: 6482: 6481:fleshy-finned 6479: 6477: 6474: 6473: 6472: 6469: 6465: 6462: 6460: 6457: 6455: 6452: 6451: 6450: 6449:Cartilaginous 6447: 6443: 6440: 6438: 6435: 6434: 6433: 6430: 6429: 6427: 6423: 6411: 6408: 6406: 6403: 6401: 6398: 6397: 6396: 6393: 6389: 6386: 6384: 6381: 6379: 6376: 6374: 6371: 6370: 6369: 6366: 6362: 6359: 6357: 6354: 6352: 6349: 6347: 6344: 6343: 6342: 6339: 6338: 6336: 6334: 6330: 6324: 6321: 6319: 6316: 6314: 6311: 6309: 6306: 6304: 6301: 6300: 6298: 6296: 6292: 6289: 6287: 6283: 6277: 6274: 6272: 6269: 6267: 6264: 6262: 6259: 6257: 6254: 6252: 6249: 6247: 6244: 6242: 6239: 6237: 6234: 6232: 6229: 6228: 6226: 6222: 6216: 6213: 6211: 6208: 6206: 6203: 6201: 6198: 6196: 6193: 6191: 6188: 6186: 6183: 6181: 6178: 6176: 6173: 6171: 6168: 6166: 6163: 6162: 6160: 6158: 6153: 6147: 6144: 6142: 6139: 6137: 6134: 6132: 6129: 6127: 6124: 6122: 6119: 6117: 6114: 6112: 6109: 6107: 6104: 6102: 6099: 6097: 6094: 6092: 6089: 6087: 6086:Electric fish 6084: 6082: 6079: 6077: 6074: 6072: 6069: 6067: 6064: 6062: 6059: 6057: 6054: 6052: 6049: 6048: 6046: 6040: 6034: 6031: 6029: 6026: 6024: 6021: 6019: 6016: 6014: 6011: 6009: 6006: 6004: 6001: 6000: 5998: 5996: 5992: 5984: 5981: 5980: 5979: 5976: 5974: 5971: 5969: 5966: 5964: 5961: 5959: 5956: 5954: 5951: 5949: 5946: 5944: 5941: 5939: 5936: 5934: 5931: 5929: 5926: 5924: 5921: 5919: 5916: 5914: 5911: 5910: 5908: 5906: 5902: 5896: 5893: 5891: 5888: 5886: 5883: 5881: 5878: 5876: 5873: 5871: 5868: 5866: 5863: 5861: 5858: 5856: 5853: 5851: 5848: 5846: 5843: 5841: 5838: 5837: 5835: 5833: 5827: 5821: 5818: 5816: 5813: 5809: 5806: 5804: 5801: 5800: 5799: 5796: 5792: 5789: 5787: 5784: 5783: 5782: 5779: 5777: 5774: 5772: 5769: 5765: 5762: 5761: 5760: 5757: 5755: 5752: 5750: 5747: 5745: 5742: 5740: 5737: 5733: 5730: 5729: 5728: 5725: 5723: 5720: 5718: 5717:Mauthner cell 5715: 5713: 5710: 5706: 5703: 5701: 5698: 5697: 5696: 5693: 5691: 5688: 5684: 5681: 5679: 5676: 5674: 5671: 5669: 5666: 5664: 5661: 5659: 5656: 5655: 5654: 5651: 5647: 5644: 5643: 5642: 5639: 5637: 5636:Chromatophore 5634: 5632: 5629: 5625: 5622: 5620: 5617: 5616: 5615: 5612: 5610: 5607: 5605: 5602: 5601: 5599: 5597: 5591: 5587: 5582: 5572: 5569: 5567: 5564: 5562: 5559: 5557: 5554: 5552: 5549: 5545: 5542: 5540: 5537: 5535: 5532: 5531: 5530: 5527: 5525: 5522: 5520: 5517: 5515: 5512: 5510: 5507: 5506: 5504: 5502: 5497: 5493: 5486: 5481: 5479: 5474: 5472: 5467: 5466: 5463: 5454: 5449: 5445: 5441: 5437: 5432: 5428: 5422: 5414: 5408: 5404: 5399: 5398: 5394: 5386: 5382: 5377: 5372: 5367: 5362: 5358: 5354: 5350: 5343: 5340: 5335: 5331: 5327: 5323: 5319: 5315: 5311: 5307: 5303: 5299: 5298: 5290: 5287: 5282: 5278: 5274: 5270: 5265: 5260: 5256: 5252: 5248: 5241: 5238: 5233: 5229: 5225: 5221: 5216: 5211: 5207: 5203: 5196: 5194: 5185: 5183: 5181: 5177: 5172: 5167:|author= 5160: 5144: 5137: 5134: 5129: 5125: 5120: 5115: 5110: 5105: 5101: 5097: 5094:(9): e44670. 5093: 5089: 5085: 5078: 5076: 5072: 5068: 5065: 5060: 5057: 5054: 5048: 5046: 5044: 5040: 5037: 5032: 5029: 5025: 5020: 5017: 5012: 5008: 5004: 5000: 4992: 4989: 4984: 4980: 4976: 4972: 4968: 4964: 4963: 4958: 4951: 4948: 4936: 4932: 4925: 4922: 4910: 4906: 4899: 4896: 4893: 4892:9780080549521 4889: 4885: 4881: 4875: 4872: 4867: 4863: 4858: 4853: 4849: 4845: 4841: 4837: 4834:: 1248–1259. 4833: 4829: 4825: 4818: 4815: 4811: 4808: 4803: 4800: 4795: 4791: 4787: 4783: 4779: 4775: 4768: 4765: 4760: 4756: 4751: 4746: 4742: 4738: 4734: 4727: 4725: 4721: 4716: 4712: 4707: 4702: 4698: 4694: 4690: 4688: 4679: 4677: 4673: 4668: 4664: 4660: 4656: 4652: 4648: 4644: 4640: 4633: 4631: 4629: 4627: 4623: 4618: 4614: 4609: 4604: 4600: 4596: 4592: 4590: 4581: 4579: 4577: 4575: 4573: 4569: 4564: 4560: 4556: 4552: 4548: 4544: 4540: 4536: 4529: 4527: 4525: 4523: 4521: 4517: 4512: 4508: 4503: 4498: 4494: 4490: 4486: 4482: 4478: 4471: 4469: 4465: 4458: 4456: 4452: 4447: 4443: 4439: 4435: 4430: 4425: 4421: 4417: 4413: 4406: 4403: 4391: 4387: 4381: 4378: 4373: 4369: 4365: 4361: 4357: 4353: 4349: 4345: 4341: 4337: 4330: 4328: 4324: 4319: 4315: 4310: 4305: 4301: 4297: 4293: 4286: 4283: 4278: 4274: 4269: 4264: 4260: 4256: 4252: 4248: 4244: 4240: 4236: 4232: 4228: 4221: 4218: 4213: 4209: 4205: 4201: 4197: 4193: 4189: 4185: 4180: 4175: 4171: 4167: 4160: 4157: 4151: 4146: 4142: 4138: 4135:(6): 060502. 4134: 4130: 4126: 4119: 4117: 4113: 4108: 4104: 4099: 4094: 4090: 4086: 4082: 4078: 4074: 4070: 4067:(1): 140163. 4066: 4062: 4058: 4051: 4048: 4040: 4034: 4031: 4026: 4022: 4018: 4014: 4010: 4006: 4002: 3998: 3994: 3987: 3985: 3981: 3976: 3972: 3968: 3964: 3960: 3956: 3952: 3948: 3944: 3940: 3932: 3930: 3926: 3921: 3917: 3913: 3909: 3905: 3901: 3897: 3893: 3889: 3885: 3878: 3876: 3872: 3859: 3852: 3849: 3844: 3840: 3835: 3830: 3826: 3822: 3818: 3814: 3810: 3806: 3802: 3795: 3792: 3787: 3783: 3778: 3773: 3769: 3765: 3760: 3755: 3751: 3747: 3743: 3736: 3734: 3730: 3725: 3721: 3716: 3711: 3707: 3703: 3700:(6): 060502. 3699: 3695: 3691: 3684: 3681: 3676: 3672: 3668: 3664: 3660: 3656: 3652: 3648: 3641: 3638: 3626: 3619: 3616: 3603: 3599: 3593: 3590: 3585: 3583:9780198549567 3579: 3575: 3571: 3565: 3563: 3561: 3557: 3552: 3545: 3542: 3539: 3533: 3531: 3529: 3527: 3523: 3518: 3517: 3509: 3506: 3501: 3494: 3491: 3487:(4): 287–305. 3486: 3482: 3475: 3472: 3460: 3456: 3449: 3446: 3441: 3435: 3431: 3430: 3422: 3419: 3415: 3409: 3406: 3390: 3383: 3377: 3374: 3369: 3365: 3360: 3355: 3351: 3347: 3343: 3339: 3335: 3331: 3327: 3320: 3318: 3314: 3309: 3305: 3301: 3297: 3293: 3289: 3285: 3281: 3274: 3272: 3268: 3263: 3259: 3255: 3251: 3244: 3242: 3240: 3236: 3231: 3227: 3222: 3217: 3213: 3209: 3205: 3201: 3197: 3193: 3189: 3182: 3179: 3174: 3170: 3165: 3160: 3156: 3152: 3148: 3141: 3139: 3135: 3130: 3126: 3122: 3118: 3114: 3110: 3106: 3102: 3098: 3094: 3087: 3084: 3079: 3075: 3070: 3065: 3061: 3057: 3052: 3047: 3043: 3039: 3035: 3028: 3025: 3021: 3019: 3011: 3008: 3004: 2998: 2995: 2990: 2986: 2981: 2976: 2971: 2966: 2962: 2958: 2954: 2950: 2946: 2939: 2936: 2931: 2927: 2922: 2917: 2913: 2909: 2908: 2907:Palaeontology 2903: 2896: 2893: 2888: 2884: 2880: 2876: 2875: 2870: 2863: 2860: 2855: 2848: 2846: 2842: 2837: 2831: 2827: 2823: 2816: 2814: 2810: 2805: 2801: 2797: 2793: 2789: 2785: 2784: 2776: 2773: 2768: 2764: 2760: 2756: 2752: 2748: 2747: 2739: 2736: 2731: 2727: 2723: 2719: 2714: 2709: 2705: 2701: 2697: 2690: 2687: 2682: 2678: 2673: 2668: 2664: 2660: 2656: 2649: 2646: 2642: 2639: 2634: 2631: 2625: 2620: 2617: 2615: 2612: 2610: 2607: 2605: 2602: 2600: 2599:Reptile scale 2597: 2595: 2592: 2590: 2587: 2585: 2582: 2580: 2577: 2576: 2572: 2570: 2568: 2563: 2562:morphological 2559: 2558: 2554: 2550: 2546: 2542: 2540: 2536: 2532: 2528: 2523: 2521: 2517: 2513: 2505: 2498: 2496: 2494: 2490: 2486: 2482: 2478: 2474: 2470: 2465: 2463: 2459: 2455: 2454:invertebrates 2450: 2447:A passage in 2442: 2436: 2429: 2424: 2420: 2413: 2408: 2406: 2404: 2400: 2395: 2392: 2385: 2379: 2374: 2370: 2364: 2359: 2351: 2346: 2342: 2335: 2331: 2330: 2324: 2320: 2316: 2313: 2312: 2311: 2309: 2305: 2301: 2297: 2293: 2285: 2281: 2278: 2275: 2271: 2268: 2265: 2261: 2258: 2254: 2251: 2250:overhand knot 2247: 2243: 2239: 2235: 2232: 2231: 2230: 2223: 2217: 2212: 2207: 2201: 2195: 2190: 2186: 2182: 2176: 2171: 2165: 2162:By contrast, 2161: 2158: 2154: 2151: 2148: 2144: 2140: 2136: 2135: 2130: 2127: 2123: 2119: 2115: 2111: 2108: 2104: 2101: 2098: 2094: 2090: 2086: 2082: 2079: 2075: 2071: 2070: 2069: 2067: 2050: 2041: 2021: 2013: 2004: 1997: 1980: 1978: 1976: 1972: 1968: 1964: 1960: 1959:triglycerides 1956: 1952: 1948: 1947:organogenesis 1944: 1939: 1937: 1933: 1929: 1924: 1923:ganoid scales 1916: 1911: 1908: 1904: 1900: 1896: 1892: 1888: 1884: 1880: 1877: 1873: 1869: 1865: 1861: 1857: 1856: 1855: 1853: 1852:metamorphosis 1849: 1848:morphogenesis 1846:occurs, then 1845: 1841: 1833: 1831: 1829: 1825: 1821: 1817: 1813: 1805: 1802: 1799: 1798: 1797: 1795: 1790: 1787: 1779: 1774: 1767: 1765: 1762: 1757: 1753: 1749: 1747: 1743: 1739: 1734: 1732: 1728: 1723: 1721: 1716: 1714: 1710: 1706: 1702: 1698: 1689: 1685: 1683: 1682: 1677: 1673: 1669: 1665: 1661: 1657: 1649: 1647: 1644: 1639: 1634: 1630: 1628: 1623: 1622:Pressure drag 1619: 1610: 1602: 1595: 1593: 1590: 1586: 1582: 1576: 1574: 1565: 1563: 1560: 1556: 1553: 1548: 1546: 1542: 1538: 1533: 1528: 1526: 1522: 1518: 1514: 1511: 1507: 1497: 1491: 1484: 1482: 1480: 1476: 1471: 1469: 1464: 1462: 1458: 1454: 1450: 1446: 1445:blood vessels 1442: 1438: 1435: 1431: 1427: 1423: 1419: 1415: 1411: 1403: 1397: 1389: 1387: 1384: 1382: 1377: 1374: 1370: 1366: 1362: 1357: 1354: 1345: 1340: 1332: 1324: 1317: 1315: 1313: 1305: 1302: 1299: 1296: 1293: 1290: 1289: 1288: 1285: 1283: 1279: 1275: 1271: 1267: 1263: 1259: 1255: 1251: 1238: 1231: 1226: 1222: 1215: 1210: 1206: 1199: 1194: 1190: 1183: 1178: 1174: 1167: 1162: 1160: 1152: 1146: 1142: 1135: 1130: 1120: 1117: 1111: 1107: 1100: 1096: 1095: 1086: 1082: 1075: 1066: 1059: 1045: 1043: 1039: 1035: 1031: 1027: 1023: 1015: 1005: 998: 993: 989: 982: 977: 973: 966: 961: 957: 950: 945: 941: 934: 929: 925: 918: 913: 909: 902: 897: 893: 886: 881: 879: 873: 864: 860: 859: 851: 842: 835: 814: 810: 809:Asian arowana 804: 800: 793: 789: 788: 785: 783: 779: 771: 769: 766: 764: 760: 756: 752: 746: 743: 740: 736: 728: 726: 724: 723:alligator gar 720: 716: 709: 706:, circa. 130 705: 704: 696: 691: 684: 679: 672: 667: 665: 663: 656: 651: 643: 639: 632: 627: 623: 619: 612: 607: 603: 599: 598:alligator gar 592: 587: 585: 583: 573: 569: 566: 565: 560: 559:Carboniferous 553: 546: 542: 541: 538: 536: 532: 528: 524: 520: 519: 514: 510: 506: 502: 498: 494: 490: 486: 482: 477: 475: 471: 470: 465: 461: 457: 453: 449: 445: 441: 434: 433:alligator gar 429: 422: 417: 411:Ganoid scales 410: 408: 406: 401: 399: 398: 393: 389: 385: 381: 380: 375: 371: 367: 366: 361: 352: 348: 343: 336: 334: 332: 328: 324: 320: 316: 312: 308: 304: 300: 296: 292: 281: 274: 272: 269: 264: 260: 257: 253: 249: 244: 242: 238: 231: 229: 224: 222: 217: 216: 210: 203: 201: 199: 195: 191: 187: 183: 179: 176:layer of the 175: 171: 166: 164: 160: 156: 152: 148: 144: 140: 136: 132: 128: 124: 120: 116: 112: 108: 104: 99: 96: 92: 88: 84: 80: 76: 72: 68: 64: 60: 51: 47: 41: 37: 33: 19: 6629:Fish anatomy 6588: 6576: 6476:spiny-finned 6425:Major groups 6146:Intelligence 6126:Scale eaters 6071:Cleaner fish 5953:Mouthbrooder 5905:Reproduction 5880:Schreckstoff 5865:Lateral line 5781:Swim bladder 5771:Spiral valve 5758: 5700:hyomandibula 5683:pseudobranch 5566:Hypoxia in - 5443: 5439: 5402: 5356: 5352: 5342: 5301: 5295: 5289: 5254: 5250: 5240: 5205: 5201: 5192: 5147:. Retrieved 5136: 5091: 5087: 5066: 5059: 5031: 5019: 5002: 4998: 4991: 4966: 4960: 4956: 4950: 4938:. Retrieved 4935:The Atlantic 4934: 4924: 4912:. Retrieved 4908: 4898: 4883: 4874: 4831: 4827: 4817: 4809: 4802: 4777: 4773: 4767: 4740: 4736: 4696: 4692: 4686: 4642: 4638: 4598: 4594: 4588: 4538: 4534: 4484: 4480: 4419: 4415: 4405: 4395:26 September 4393:. Retrieved 4389: 4380: 4339: 4335: 4302:(4): 36–44. 4299: 4295: 4285: 4234: 4230: 4220: 4169: 4165: 4159: 4132: 4128: 4064: 4060: 4050: 4033: 4000: 3996: 3942: 3938: 3887: 3883: 3862:. Retrieved 3851: 3808: 3804: 3794: 3749: 3745: 3697: 3693: 3683: 3650: 3646: 3640: 3628:. Retrieved 3618: 3606:. Retrieved 3601: 3592: 3573: 3550: 3544: 3515: 3508: 3499: 3493: 3484: 3480: 3474: 3462:. Retrieved 3458: 3448: 3428: 3421: 3413: 3408: 3396:. Retrieved 3389:the original 3376: 3333: 3329: 3283: 3279: 3253: 3249: 3195: 3191: 3181: 3157:(1): 86–95. 3154: 3150: 3096: 3092: 3086: 3041: 3037: 3027: 3022:55: 459–470. 3017: 3010: 2997: 2952: 2948: 2938: 2911: 2905: 2895: 2878: 2872: 2868: 2862: 2853: 2821: 2787: 2781: 2775: 2750: 2744: 2738: 2703: 2699: 2689: 2662: 2658: 2648: 2640: 2633: 2614:Snake scales 2609:Scale armour 2555: 2543: 2524: 2516:scale-eating 2515: 2510: 2468: 2466: 2458:abominations 2446: 2396: 2393: 2390: 2308:sticklebacks 2289: 2270:Mandarinfish 2234:Jawless fish 2228: 2222:Mandarinfish 2167:vertebrates. 2141:between the 2138: 2132: 2125: 2074:lateral line 2063: 2003:common roach 1988:Lateral line 1940: 1920: 1895:invagination 1837: 1828:lateral line 1812:pineconefish 1809: 1793: 1791: 1784: 1778:Pineconefish 1754: 1750: 1746:lotus effect 1735: 1724: 1717: 1709:green sludge 1694: 1679: 1653: 1635: 1631: 1627:laminar flow 1615: 1577: 1572: 1569: 1549: 1545:laminar flow 1543:forcing the 1529: 1525:lotus effect 1521:anti-fouling 1510:hydrodynamic 1508:that reduce 1502: 1475:ostracoderms 1472: 1465: 1457:vitrodentine 1425: 1409: 1408: 1385: 1378: 1376:camouflage. 1358: 1353:transparency 1349: 1309: 1304:True ctenoid 1303: 1297: 1291: 1286: 1268:between the 1262:lateral line 1260:, along the 1247: 1118: 1025: 1019: 856: 775: 767: 763:age the fish 758: 754: 747: 732: 713: 701: 659: 579: 562: 551: 535:tooth enamel 516: 478: 474:vitrodentine 472:in place of 467: 444:paddlefishes 437: 402: 395: 377: 363: 357: 327:vitrodentine 288: 265: 261: 245: 234: 226: 219: 213: 167: 123:ostracoderms 115:anglerfishes 103:shrimpfishes 100: 86: 67:jawed fishes 58: 56: 46:teleost fish 36: 6590:WikiProject 6549:Prehistoric 6533:Threatened 6224:Other types 6121:Sardine run 6096:Forage fish 6076:Corallivory 5928:Development 5913:Bubble nest 5786:physoclisti 5776:Suckermouth 5749:Root effect 5571:Ichthyology 5353:BMC Biology 4687:Danio rerio 4589:Danio rerio 3746:Biomimetics 3464:29 December 3414:Ichthyology 2881:(1): 1–11. 2869:Phlebolepis 2512:Lepidophagy 2499:Lepidophagy 2319:exoskeleton 2143:pelvic fins 2103:Surgeonfish 2032:Surgeonfish 1975:interaction 1963:cholesterol 1928:osteoblasts 1909:will arise. 1899:basal cells 1876:fibroblasts 1654:The rough, 1552:collagenous 1515:and reduce 1496:tiger shark 1441:pulp cavity 1266:fibroblasts 518:Cheirolepis 505:coelacanths 421:spotted gar 392:coelacanths 241:fossil fish 215:Paralogania 184:. The same 127:bony fishes 83:colouration 6205:Groundfish 6200:Freshwater 6195:Euryhaline 6180:Coral reef 6116:Salmon run 6106:Paedophagy 6008:Amphibious 5995:Locomotion 5803:pharyngeal 5791:physostome 5744:Photophore 5690:Glossohyal 5663:gill raker 5646:dorsal fin 5596:physiology 5145:. FishBase 4940:26 January 4179:1904.12752 2641:Etymonline 2626:References 2280:Anglerfish 2274:aposematic 2200:Pufferfish 2164:pufferfish 1907:primordium 1903:epithelium 1844:mesenchyme 1820:threadfins 1589:biomimetic 1517:turbulence 1485:Shark skin 1479:placoderms 1434:vertebrate 1430:homologous 1318:Reflection 1312:flatfishes 1258:caudal fin 1254:integument 1145:Crazy fish 1085:perch-like 1030:perch-like 757:(singular 737:(the more 622:scute-like 602:rhomboidal 582:rhomboidal 531:homologous 527:acanthodii 513:stem group 495:, such as 487:bones and 388:lungfishes 370:coelacanth 360:imbricated 351:coelacanth 307:coelacanth 301:), and in 297:(subclass 295:lungfishes 268:convergent 237:thelodonts 194:homologous 119:morphology 91:Old French 79:reflection 75:camouflage 59:fish scale 6454:chimaeras 6341:Predatory 6318:Salmonids 6276:Whitefish 6266:Poisonous 6241:Diversity 6175:Coldwater 6111:Predatory 6101:Migratory 6061:Bait ball 6044:behaviour 5963:Pregnancy 5958:Polyandry 5732:papillare 5727:Operculum 5722:Meristics 5668:gill slit 5631:Cleithrum 5561:Fish kill 5551:Fear of - 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