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Cycloid scales cover these teleost fish (rohu)

A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as well as possible hydrodynamic advantages. The term scale derives from the Old French "escale", meaning a shell pod or husk.[1]

Scales vary enormously in size, shape, structure, and extent, ranging from strong and rigid armour plates in fishes such as shrimpfishes and boxfishes, to microscopic or absent in fishes such as eels and anglerfishes. The morphology of a scale can be used to identify the species of fish it came from.

Most bony fishes are covered with the cycloid scales of salmon and carp, or the ctenoid scales of perch, or the ganoid scales of sturgeons and gars. Cartilaginous fishes (sharks and rays) are covered with placoid scales. Some species are covered instead by scutes, and others have no outer covering on part or all of the skin.

Fish scales are part of the fish's integumentary system, and are produced from the mesoderm layer of the dermis, which distinguishes them from reptile scales.[2] The same genes involved in tooth and hair development in mammals are also involved in scale development. The placoid scales of cartilaginous fishes are also called dermal denticles and are structurally homologous with vertebrate teeth. It has been suggested that the scales of bony fishes are similar in structure to teeth, but they probably originate from different tissue.[3] Most fish are also covered in a protective layer of mucus (slime).


Thelodont scalesEdit

Left to right: denticles of Paralogania (?), Shielia taiti, Lanarkia horrida

The bony scales of thelodonts, the most abundant form of fossil fish, are well understood. The scales were formed and shed throughout the organisms' lifetimes, and quickly separated after their death.[4]

Bone, a tissue that is both resistant to mechanical damage and relatively prone to fossilization, often preserves internal detail, which allows the histology and growth of the scales to be studied in detail. The scales comprise a non-growing "crown" composed of dentine, with a sometimes-ornamented enameloid upper surface and an aspidine base.[5] Its growing base is made of cell-free bone, which sometimes developed anchorage structures to fix it in the side of the fish.[6] Beyond that, there appear to be five types of bone-growth, which may represent five natural groupings within the thelodonts—or a spectrum ranging between the end members meta- (or ortho-) dentine and mesodentine tissues.[7] 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.[6]

However, using scale morphology alone to distinguish species has some pitfalls. Within each organism, scale shape varies hugely according to body area,[8] with intermediate forms appearing between different areas—and to make matters worse, scale morphology may not even be constant within one area. To confuse things further, scale morphologies are not unique to taxa, and may be indistinguishable on the same area of two different species.[9]

The morphology and histology of thelodonts provides the main tool for quantifying their diversity and distinguishing between species, although ultimately using such convergent traits is prone to errors. Nonetheless, a framework comprising three groups has been proposed based upon scale morphology and histology.[7] Comparisons to modern shark species have shown that thelodont scales were functionally similar to those of modern cartilaginous fish, and likewise has allowed an extensive comparison between ecological niches.[10]

Cosmoid scalesEdit

Queensland lungfish

True cosmoid scales are not found on extant fish. They are found only on ancient lobe-finned fishes, including some of the earliest lungfishes. They were probably derived from a fusion of placoid scales. The inner part of the scales is made of dense lamellar bone called isopedine. On top of this lies a layer of spongy or vascular bone supplied with blood vessels, followed by a complex dentine-like layer called cosmine with a superficial outer coating of vitrodentine. The upper surface is keratin. Cosmoid scales increase in size through the growth of the lamellar bone layer.

Elasmoid scalesEdit

Lobe-finned fishes, like this preserved coelacanth, have elasmoid scales

Elasmoid scales are thin, imbricated scales composed of a layer of dense, lamellar bone called isopedine, above which is a layer of tubercles usually composed of bone, as in Eusthenopteron. The layer of dentine that was present in the first lobe-finned fish is usually reduced, as in the extant coelacanth, or entirely absent, as in extant lungfish and in the Devonian Eusthenopteron.[11] Elasmoid scales have appeared several times over the course of fish evolution. They are present in some lobe-finned fishes, such as all extant and some extinct lungfishes, as well as the coelacanths which have modified cosmoid scales that lack cosmine and are thinner than true cosmoid scales. They are also present in some tetrapodomorphs like Eusthenopteron, amiids, and teleosts, whose cycloid and ctenoid scales represent the least mineralized elasmoid scales.

Ganoid scalesEdit

Ganoid scales are found in the sturgeons, paddlefishes, gars, bowfin, and bichirs. They are derived from cosmoid scales and often have serrated edges. They are covered with a layer of hard enamel-like dentine in the place of cosmine, and a layer of inorganic bone salt called ganoine in place of vitrodentine.


Amblypterus striatus
Ganoid scales of the extinct Carboniferous fish, Amblypterus striatus. (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.  

Most ganoid scales are rhomboidal or 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.[12] In this way, ganoid scales are nearly impenetrable and are excellent protection against predation.

The ganoid scales on a bowfin are reduced in size and resemble cycloid scales
Geometrically laid out ganoid scales on a bichir

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 cycloid scales.

Native Americans and people of the Caribbean used the tough ganoid scales of the alligator gar for arrow heads, breastplates, and as shielding to cover plows. In current times jewellery is made from these scales.[13]

Leptoid scalesEdit

Leptoid (bony-ridge) scales are found on higher-order bony fish, the teleosts (the more derived clade 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, further scales are added in concentric layers as the fish grows.[14]

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 drag.[15] The scales of some species exhibit bands of uneven seasonal growth called annuli (singular annulus). These bands can be used to age the fish.

Leptoid scales come in two forms: cycloid and ctenoid.

Cycloid scalesEdit

Cycloid (circular) scales
The cycloid scale of a carp has a smooth outer edge
This Poropuntius huguenini is a carp-like fish with circular cycloid scales that are smooth to the touch

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 salmon and carp.

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

Ctenoid scalesEdit

Ctenoid (toothed) scales
The ctenoid scale of a perch has a toothed outer edge (at top of image)
This dottyback is a perch-like fish with toothed ctenoid scales that are rough to the touch
Three ctenoid scales from various locations of a perch were stained. Significant variation can be observed between the medial (middle of the fish), dorsal (top), and caudal (tail end) scales. The ctentii of each of the scales is labeled.

Ctenoid (toothed) scales are like cycloid scales, with small teeth along their outer edges. They are usually found on fishes with spiny fin rays, such as the perch-like fishes. The scales have a rough texture with a toothed outer or posterior edge featuring tiny teeth called ctenii. These scales contain almost no bone, being composed of a surface layer containing hydroxyapatite and calcium carbonate and a deeper layer composed mostly of collagen. The enamel of the other scale types is reduced to superficial ridges and ctenii.

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

Ctenoid scales, similar to other epidermal structures, originate from placodes and distinctive cellular differentiation makes them exclusive from other structures that arise from the integument.[16] Development starts near the caudal fin, along the lateral line of the fish.[17] The development process begins with an accumulation of fibroblasts between the epidermis and dermis.[16] Collagen fibrils begin to organize themselves in the dermal layer, which leads to the initiation of mineralization.[16] The circumference of the scales grows first, followed by thickness when overlapping layers mineralize together.[16]

Ctenoid scales can be further subdivided into three types:

  • Crenate scales, where the margin of the scale bears indentations and projections.
  • Spinoid scales, where the scale bears spines that are continuous with the scale itself.
  • True ctenoid scales, where the spines on the scale are distinct structures.

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


The herring's reflectors are nearly vertical for camouflage from the side.
The deep sea hatchetfish has scales which reflect blue light
The scales of a typical teleost fish, like this Atlantic herring, are silvered

Many teleost fish are covered with highly reflective scales, giving 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 transparency 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.[18]

The marine hatchetfish is extremely flattened laterally (side to side), leaving the body just millimetres thick, and the body is so silvery as to resemble aluminium foil. The mirrors consist of microscopic structures similar to those used to provide structural coloration: stacks of between 5 and 10 crystals of guanine 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 camouflage.[18]

Most fish in the upper ocean are camouflaged by silvering. In fish such as the herring, 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.[18]

Fish scales with these properties are used in some cosmetics, since they can give a shimmering effect to makeup and lipstick.[19]

Placoid scalesEdit

Cartilaginous fishes, like this tiger shark, have placoid scales (dermal denticles)
Placoid scales as viewed through an electron microscope. Also called dermal denticles, these are structurally homologous with vertebrate teeth.

Placoid (pointed, tooth-shaped) scales are found in the cartilaginous fishes: sharks, rays, and chimaeras. They are also called dermal denticles. Placoid scales are structurally homologous with vertebrate teeth ("denticle" translates to "small tooth"), having a central pulp cavity supplied with blood vessels, surrounded by a conical layer of dentine, all of which sits on top of a rectangular basal plate that rests on the dermis. The outermost layer is composed of vitrodentine, a largely inorganic enamel-like substance. Placoid scales cannot grow in size, but rather more scales are added as the fish increases in size.

Similar scales can also be found under the head of the denticle herring. The amount of scale coverage is much less in rays and chimaeras.

Shark skinEdit

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 vortices that reduce hydrodynamic drag, and reduce the turbulence, making swimming both more efficient, and quieter, compared to that of bony fishes.[20] It also serves a role in anti-fouling by exhibiting the lotus effect.[21]

Unlike bony fish, sharks have a complicated dermal corset made of flexible collagenous fibers arranged as a helical network surrounding their body. The corset works as an outer skeleton, providing attachment for their swimming muscles and thus saving energy.[22] 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 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.[23]

Scale morphologyEdit

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 stratum laxum.  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. These riblets serve a major hydrodynamic role and have shown to reduce drag by up to 9% in biomimetic test specimens. The spacing between these riblets and their height has been the subject of numerous experiments and has been a research topic. This spacing and height is consistent in the fast swimming sharks[24]

Drag reductionEdit

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 pushes the vortex further up from the surface, interacting only with the riblet tips, not causing any high-veloctiy 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.[24]

The rough, sandpaper-like texture of shark and ray skin, coupled with its toughness, has led it to be valued as a source of rawhide leather, called shagreen. One of the many historical applications of shark shagreen was in making hand-grips for swords. The rough texture of the skin is also used in Japanese cuisine to make graters called oroshiki, by attaching pieces of shark skin to wooden boards. The small size of the scales grates the food very finely.

Technical applicationEdit

There are many examples of biomimetic materials 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.

Surfaces that mimic the skin of sharks have also been used in order to keep microorganisms and algae from coating the hulls of submarines and ships. One variety is traded as "sharklet".[25][26]

A lot of the new methods for replicating shark skin involve the use of polydimethylsiloxane (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 superhydrophobic properties, exhibiting the lotus effect.[25] One study found that these biomimetic surfaces reduced drag by up to 9%,[23] while with flapping motion drag reduction reached 12.3%.[27]


Pineconefish are covered in scutes

A scute is another, less common, type of scale. Scute comes from Latin for shield, and can take the form of:

  • an external shield-like bony plate, or
  • a modified, thickened scale that often is keeled or spiny, or
  • 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.

Some fish, such as pineconefish, are completely or partially covered in scutes. River herrings and threadfins have an abdominal row of scutes, which are scales with raised, sharp points that are used for protection. Some jacks have a row of scutes following the lateral line on either side.


The cycloid scales of a common roach. The series of lateral line scales is visible in the lower half of the image.

Different groups of fish have evolved a number of modified scales to serve various functions.

Many groups of bony fishes, including pipefishes and seahorses, several families of catfishes, sticklebacks, and poachers, have developed external bony plates, structurally resembling placoid scales, as protective armour. In the boxfishes, the plates are all fused together to form a rigid shell enclosing the entire body. Yet these bony plates are not modified scales, but skin that has been ossified.

The size of the teeth on ctenoid scales can vary with position, as these scales from the rattail Cetonurus crassiceps show
Eels seem scaleless, yet some species are covered with tiny smooth cycloid scales

See alsoEdit


  1. ^ Scale Etymonline. Retrieved 28 April 2019.
  2. ^ Sharpe, P. T. (2001). "Fish scale development: Hair today, teeth and scales yesterday?". Current Biology. 11 (18): R751–R752. doi:10.1016/S0960-9822(01)00438-9. PMID 11566120.
  3. ^ Perkins, Sid (16 October 2013). "The First False Teeth". Science. Retrieved 2 March 2018.
  4. ^ Turner, S.; Tarling, D. H. (1982). "Thelodont and other agnathan distributions as tests of Lower Paleozoic continental reconstructions". Palaeogeography, Palaeoclimatology, Palaeoecology. 39 (3–4): 295–311. doi:10.1016/0031-0182(82)90027-X.
  5. ^ Märss, T. (2006). "Exoskeletal ultrasculpture of early vertebrates". Journal of Vertebrate Paleontology. 26 (2): 235–252. doi:10.1671/0272-4634(2006)26[235:EUOEV]2.0.CO;2.
  6. ^ a b Janvier, Philippe (1998). "Early vertebrates and their extant relatives". Early Vertebrates. Oxford University Press. pp. 123–127. ISBN 978-0-19-854047-2.
  7. ^ a b Turner, S. (1991). "Monophyly and interrelationships of the Thelodonti". In M. M. Chang, Y. H. Liu & G. R. Zhang (eds.). Early Vertebrates and Related Problems of Evolutionary Biology. Science Press, Beijing. pp. 87–119.CS1 maint: Uses editors parameter (link)
  8. ^ Märss, T. (1986). "Squamation of the thelodont agnathan Phlebolepis". Journal of Vertebrate Paleontology. 6 (1): 1–11. doi:10.1080/02724634.1986.10011593.
  9. ^ Botella, H.; J. I. Valenzuela-Rios; P. Carls (2006). "A New Early Devonian thelodont from Celtiberia (Spain), with a revision of Spanish thelodonts". Palaeontology. 49 (1): 141–154. doi:10.1111/j.1475-4983.2005.00534.x.
  10. ^ Ferrón, Humberto G.; Botella, Héctor (2017). "Squamation and ecology of thelodonts". PLoS ONE. 12 (2): e0172781. doi:10.1371/journal.pone.0172781. PMC 5328365. PMID 28241029.
  11. ^ Zylberberg, L., Meunier, F.J., Laurin, M. (2010). A microanatomical and histological study of the postcranial dermal skeleton in the Devonian sarcopterygian Eusthenopteron foordi, Acta Palaeontologica Polonica 55: 459–470.
  12. ^ Sherman, Vincent R.; Yaraghi, Nicholas A.; Kisailus, David; Meyers, Marc A. (2016-12-01). "Microstructural and geometric influences in the protective scales of Atractosteus spatula". Journal of the Royal Society Interface. 13 (125): 20160595. doi:10.1098/rsif.2016.0595. ISSN 1742-5689. PMC 5221522. PMID 27974575.
  13. ^ "Missouri Alligator Gar Management and Restoration Plan" (PDF). Missouri Department of Conservation Fisheries Division. January 22, 2013. Archived from the original (PDF) on May 6, 2016. Retrieved April 12, 2019.
  14. ^ Lagler, K. F., J. E. Bardach, and R. R. Miller (1962) Ichthyology. New York: John Wiley & Sons.
  15. ^ Ballard, Bonnie; Cheek, Ryan (2 July 2016). Exotic Animal Medicine for the Veterinary Technician. John Wiley & Sons. ISBN 978-1-118-92421-1.
  16. ^ a b c d Kawasaki, Kenta C., "A Genetic Analysis of Cichlid Scale Morphology" (2016). Masters Theses May 2014 - current. 425.
  17. ^ Helfman, Gene (2009). The Diversity of Fishes Biology, Evolution, and Ecology. Wiley-Blackwell.
  18. ^ a b c Herring, Peter (2002). The Biology of the Deep Ocean. Oxford: Oxford University Press. pp. 193–195. ISBN 9780198549567.
  19. ^ "There Are Probably Fish Scales In Your Lipstick". HuffPost India. 2015-04-23. Retrieved 2019-05-06.
  20. ^ Martin, R. Aidan. "Skin of the Teeth". Retrieved 2007-08-28.
  21. ^ Fürstner, Reiner; Barthlott, Wilhelm; Neinhuis, Christoph; Walzel, Peter (2005-02-01). "Wetting and Self-Cleaning Properties of Artificial Superhydrophobic Surfaces". Langmuir. 21 (3): 956–961. doi:10.1021/la0401011. ISSN 0743-7463. PMID 15667174.
  22. ^ Martin, R. Aidan. "The Importance of Being Cartilaginous". ReefQuest Centre for Shark Research. Retrieved 2009-08-29.
  23. ^ a b Hage, W.; Bruse, M.; Bechert, D. W. (2000-05-01). "Experiments with three-dimensional riblets as an idealized model of shark skin". Experiments in Fluids. 28 (5): 403–412. doi:10.1007/s003480050400. ISSN 1432-1114.
  24. ^ a b Motta, Philip; Habegger, Maria Laura; Lang, Amy; Hueter, Robert; Davis, Jessica (2012-10-01). "Scale morphology and flexibility in the shortfin mako Isurus oxyrinchus and the blacktip shark Carcharhinus limbatus". Journal of Morphology. 273 (10): 1096–1110. doi:10.1002/jmor.20047. ISSN 1097-4687. PMID 22730019.
  25. ^ a b Liu, Yunhong; Li, Guangji (2012-12-15). "A new method for producing "Lotus Effect" on a biomimetic shark skin". Journal of Colloid and Interface Science. 388 (1): 235–242. doi:10.1016/j.jcis.2012.08.033. ISSN 0021-9797. PMID 22995249.
  26. ^ "Sharklet Discovery | Sharklet Technologies, Inc". Retrieved 2018-09-26.
  27. ^ Lauder, George V.; Oeffner, Johannes (2012-03-01). "The hydrodynamic function of shark skin and two biomimetic applications". Journal of Experimental Biology. 215 (5): 785–795. doi:10.1242/jeb.063040. ISSN 1477-9145. PMID 22323201.

Further readingEdit

  • Helfman, G.S., B.B. Collette and D.E. Facey (1997). The Diversity of Fishes. Blackwell Science. pp. 33–36. ISBN 978-0-86542-256-8.CS1 maint: Multiple names: authors list (link)

External linksEdit

  • Hydrodynamic aspects of shark scales [1]
  • Fish scales and flow manipulation [2]