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Countershading

From Wikipedia, in a visual modern way
Many animals, such as this grey reef shark, are countershaded.
Many animals, such as this grey reef shark, are countershaded.
Illustration from the artist Abbot Thayer's  1909 book on camouflage of a Luna caterpillar Actias lunaa) in position b) inverted.
Illustration from the artist Abbot Thayer's 1909 book on camouflage of a Luna caterpillar Actias luna
a) in position b) inverted.

Countershading, or Thayer's law, is a method of camouflage in which an animal's coloration is darker on the top or upper side and lighter on the underside of the body.[1] This pattern is found in many species of mammals, reptiles, birds, fish, and insects, both in predators and in prey.

When light falls from above on a uniformly coloured three-dimensional object such as a sphere, it makes the upper side appear lighter and the underside darker, grading from one to the other. This pattern of light and shade makes the object appear solid, and therefore easier to detect. The classical form of countershading, discovered in 1909 by the artist Abbott Handerson Thayer, works by counterbalancing the effects of self-shadowing, again typically with grading from dark to light. In theory this could be useful for military camouflage, but in practice it has rarely been applied, despite the best efforts of Thayer and, later, in the Second World War, of the zoologist Hugh Cott.

The precise function of various patterns of animal coloration that have been called countershading has been debated by zoologists such as Hannah Rowland (2009), with the suggestion that there may be multiple functions including flattening and background matching when viewed from the side; background matching when viewed from above or below, implying separate colour schemes for the top and bottom surfaces; outline obliteration from above; and a variety of other largely untested non-camouflage theories. A related mechanism, counter-illumination, adds the creation of light by bioluminescence or lamps to match the actual brightness of a background. Counter-illumination camouflage is common in marine organisms such as squid. It has been studied up to the prototype stage for military use in ships and aircraft, but it too has rarely or never been used in warfare.

The reverse of countershading, with the belly pigmented darker than the back, enhances contrast and so makes animals more conspicuous. It is found in animals that can defend themselves, such as skunks. The pattern is used both in startle or deimatic displays and as a signal to warn off experienced predators. However, animals that habitually live upside-down but lack strong defences, such as the Nile catfish and the Luna moth caterpillar, have upside-down countershading for camouflage.

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Camouflage

Camouflage

Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the battledress of a modern soldier, and the leaf-mimic katydid's wings. A third approach, motion dazzle, confuses the observer with a conspicuous pattern, making the object visible but momentarily harder to locate, as well as making general aiming easier. The majority of camouflage methods aim for crypsis, often through a general resemblance to the background, high contrast disruptive coloration, eliminating shadow, and countershading. In the open ocean, where there is no background, the principal methods of camouflage are transparency, silvering, and countershading, while the ability to produce light is among other things used for counter-illumination on the undersides of cephalopods such as squid. Some animals, such as chameleons and octopuses, are capable of actively changing their skin pattern and colours, whether for camouflage or for signalling. It is possible that some plants use camouflage to evade being eaten by herbivores.

Animal coloration

Animal coloration

Animal coloration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male has strong patterns, conspicuous colours and is iridescent, while the female is far less visible.

Mammal

Mammal

A mammal is a vertebrate animal of the class Mammalia. Mammals are characterized by the presence of milk-producing mammary glands for feeding their young, a neocortex region of the brain, fur or hair, and three middle ear bones. These characteristics distinguish them from reptiles and birds, which they diverged from in the Carboniferous Period over 300 million years ago. Around 6,400 extant species of mammals have been described and divided into 29 orders.

Bird

Bird

Birds are a group of warm-blooded vertebrates constituting the class Aves, characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweight skeleton. Birds live worldwide and range in size from the 5.5 cm (2.2 in) bee hummingbird to the 2.8 m common ostrich. There are about ten thousand living species, more than half of which are passerine, or "perching" birds. Birds have wings whose development varies according to species; the only known groups without wings are the extinct moa and elephant birds. Wings, which are modified forelimbs, gave birds the ability to fly, although further evolution has led to the loss of flight in some birds, including ratites, penguins, and diverse endemic island species. The digestive and respiratory systems of birds are also uniquely adapted for flight. Some bird species of aquatic environments, particularly seabirds and some waterbirds, have further evolved for swimming.

Fish

Fish

Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of living fish species are ray-finned fish, belonging to the class Actinopterygii, with around 99% of those being teleosts.

Insect

Insect

Insects are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body, three pairs of jointed legs, compound eyes and one pair of antennae. Their blood is not totally contained in vessels; some circulates in an open cavity known as the haemocoel. Insects are the most diverse group of animals; they include more than a million described species and represent more than half of all known living organisms. The total number of extant species is estimated at between six and ten million; potentially over 90% of the animal life forms on Earth are insects. Insects may be found in nearly all environments, although only a small number of species reside in the oceans, which are dominated by another arthropod group, crustaceans, which recent research has indicated insects are nested within.

Abbott Handerson Thayer

Abbott Handerson Thayer

Abbott Handerson Thayer was an American artist, naturalist and teacher. As a painter of portraits, figures, animals and landscapes, he enjoyed a certain prominence during his lifetime, and his paintings are represented in the major American art collections. He is perhaps best known for his 'angel' paintings, some of which use his children as models.

Counter-illumination

Counter-illumination

Counter-illumination is a method of active camouflage seen in marine animals such as firefly squid and midshipman fish, and in military prototypes, producing light to match their backgrounds in both brightness and wavelength.

Bioluminescence

Bioluminescence

Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some bioluminescent bacteria, and terrestrial arthropods such as fireflies. In some animals, the light is bacteriogenic, produced by symbiotic bacteria such as those from the genus Vibrio; in others, it is autogenic, produced by the animals themselves.

Deimatic behaviour

Deimatic behaviour

Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey animal an opportunity to escape. The term deimatic or dymantic originates from the Greek δειματόω (deimatóo), meaning "to frighten".

Aposematism

Aposematism

Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or smell, sharp spines, or aggressive nature. These advertising signals may take the form of conspicuous coloration, sounds, odours, or other perceivable characteristics. Aposematic signals are beneficial for both predator and prey, since both avoid potential harm.

Luna moth

Luna moth

The Luna moth also known as the American moon moth is a Nearctic moth in the family Saturniidae, subfamily Saturniinae, a group commonly known as giant silk moths. It has lime-green colored wings and a white body. The larvae (caterpillars) are also green. Typically, it has a wingspan of roughly 114 mm (4.5 in), but can exceed 178 mm (7.0 in), making it one of the larger moths in North America. Across Canada, it has one generation per year, with the winged adults appearing in late May or early June, whereas farther south it will have two or even three generations per year, the first appearance as early as March in southern parts of the United States.

Early research

Thayer's 1902 patent application. He failed to convince the US Navy.
Thayer's 1902 patent application. He failed to convince the US Navy.

The English zoologist Edward Bagnall Poulton, author of The Colours of Animals (1890) discovered the countershading of various insects, including the pupa or chrysalis of the purple emperor butterfly, Apatura iris,[2] the caterpillar larvae of the brimstone moth, Opisthograptis luteolata [a] and of the peppered moth, Biston betularia.[b][3][4] However he did not use the term countershading, nor did he suggest that the effect occurred widely.[5]

Thayer's "White fowl, lacking counter-shading, against a flat white cloth."
Thayer's "White fowl, lacking counter-shading, against a flat white cloth."
A 1917 photograph of a countershading study by Thayer, who became obsessed by the mistaken idea that all animals are countershaded.[6]
A 1917 photograph of a countershading study by Thayer, who became obsessed by the mistaken idea that all animals are countershaded.[6]

The New Hampshire artist Abbott Handerson Thayer was one of the first to study and write about countershading. In his 1909 book Concealing-Coloration in the Animal Kingdom, he correctly described and illustrated countershading with photographs and paintings, but wrongly claimed that almost all animals are countershaded.[7] For this reason countershading is sometimes called Thayer's law. Thayer wrote:

Animals are painted by Nature darkest on those parts which tend to be most lighted by the sky's light, and vice versa. ... the fact that a vast majority of creatures of the whole animal kingdom wear this gradation, developed to an exquisitely minute degree, and are famous for being hard to see in their homes, speaks for itself.

— Thayer[8]

Thayer observed and painted a number of examples, including the Luna moth caterpillar Actias luna, both in its habitual upside-down feeding position, where its countershading makes it appear flat, and artificially inverted from that position, where sunlight and its inverted countershading combine to make it appear heavily shaded and therefore solid.[9] Thayer obtained a patent in 1902 to paint warships, both submarines and surface ships, using countershading,[10] but failed to convince the US Navy to adopt his ideas.[11]

Hugh Bamford Cott in his 1940 book Adaptive Coloration in Animals described many instances of countershading, following Thayer in general approach[12] but criticising Thayer's excessive claim ("He says 'All patterns and colors whatsoever of all animals that ever prey or are preyed upon are under certain normal circumstances obliterative.'") that effectively all animals are camouflaged with countershading. Cott called this "Thayer straining the theory to a fantastic extreme".[13]

Both Thayer and Cott included in their books photographs of a non-countershaded white cockerel against a white background, to make the point that in Thayer's words "a monochrome object can not be 'obliterated', no matter what its background"[14] or in Cott's words "Colour resemblance alone is not sufficient to afford concealment".[15] Cott explained that

Contrary to what might have been expected by any one lacking in artistic perception, the bird appears highly conspicuous, the back looking lighter, and the breast darker, than the background, although in actual fact, back, background and breast are all pure white."[16]

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England

England

England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea area of the Atlantic Ocean to the southwest. It is separated from continental Europe by the North Sea to the east and the English Channel to the south. The country covers five-eighths of the island of Great Britain, which lies in the North Atlantic, and includes over 100 smaller islands, such as the Isles of Scilly and the Isle of Wight.

Edward Bagnall Poulton

Edward Bagnall Poulton

Sir Edward Bagnall Poulton, FRS HFRSE FLS was a British evolutionary biologist, a lifelong advocate of natural selection through a period in which many scientists such as Reginald Punnett doubted its importance. He invented the term sympatric for evolution of species in the same place, and in his book The Colours of Animals (1890) was the first to recognise frequency-dependent selection. Poulton is also remembered for his pioneering work on animal coloration. He is credited with inventing the term aposematism for warning coloration, as well as for his experiments on 'protective coloration' (camouflage). Poulton became Hope Professor of Zoology at the University of Oxford in 1893.

The Colours of Animals

The Colours of Animals

The Colours of Animals is a zoology book written in 1890 by Sir Edward Bagnall Poulton (1856–1943). It was the first substantial textbook to argue the case for Darwinian selection applying to all aspects of animal coloration. The book also pioneered the concept of frequency-dependent selection and introduced the term "aposematism".

Apatura iris

Apatura iris

Apatura iris, the purple emperor, is a Palearctic butterfly of the family Nymphalidae.

New Hampshire

New Hampshire

New Hampshire is a state in the New England region of the Northeastern United States. It is bordered by Massachusetts to the south, Vermont to the west, Maine and the Gulf of Maine to the east, and the Canadian province of Quebec to the north. Of the 50 U.S. states, New Hampshire is the fifth smallest by area and the tenth least populous, with slightly more than 1.3 million residents as of the 2020 census. Concord is the state capital, while Manchester is the largest city. New Hampshire's motto, "Live Free or Die", reflects its role in the American Revolutionary War; its nickname, "The Granite State", refers to its extensive granite formations and quarries. It is well known nationwide for holding the first primary in the U.S. presidential election cycle, and for its resulting influence on American electoral politics.

Abbott Handerson Thayer

Abbott Handerson Thayer

Abbott Handerson Thayer was an American artist, naturalist and teacher. As a painter of portraits, figures, animals and landscapes, he enjoyed a certain prominence during his lifetime, and his paintings are represented in the major American art collections. He is perhaps best known for his 'angel' paintings, some of which use his children as models.

Concealing-Coloration in the Animal Kingdom

Concealing-Coloration in the Animal Kingdom

Concealing-Coloration in the Animal Kingdom: An Exposition of the Laws of Disguise Through Color and Pattern; Being a Summary of Abbott H. Thayer’s Discoveries is a book published ostensibly by Gerald H. Thayer in 1909, and revised in 1918, but in fact a collaboration with and completion of his father Abbott Handerson Thayer's major work.

Submarine

Submarine

A submarine is a watercraft capable of independent operation underwater. It differs from a submersible, which has more limited underwater capability. The term is also sometimes used historically or colloquially to refer to remotely operated vehicles and robots, as well as medium-sized or smaller vessels, such as the midget submarine and the wet sub. Submarines are referred to as boats rather than ships irrespective of their size.

Warship

Warship

A warship or combatant ship is a naval ship that is built and primarily intended for naval warfare. Usually they belong to the armed forces of a state. As well as being armed, warships are designed to withstand damage and are typically faster and more maneuverable than merchant ships. Unlike a merchant ship, which carries cargo, a warship typically carries only weapons, ammunition and supplies for its crew. Warships usually belong to a navy, though they have also been operated by individuals, cooperatives and corporations.

Adaptive Coloration in Animals

Adaptive Coloration in Animals

Adaptive Coloration in Animals is a 500-page textbook about camouflage, warning coloration and mimicry by the Cambridge zoologist Hugh Cott, first published during the Second World War in 1940; the book sold widely and made him famous.

Application

In animals

Ibexes are effectively flattened by countershading, making them nearly invisible against a desert background. There are three in the image.
Ibexes are effectively flattened by countershading, making them nearly invisible against a desert background. There are three in the image.

Countershading is observed in a wide range of animal groups, both terrestrial, such as deer, and marine, such as sharks.[17] It is the basis of camouflage in both predators and prey.[18] It is used alongside other forms of camouflage including colour matching and disruptive coloration.[18] Among predatory fish, the gray snapper, Lutianus griseus, is effectively flattened by its countershading, while it hunts an "almost invisible" prey, the hardhead fish, Atherina laticeps which swims over greyish sands.[19] Other countershaded marine animals include blue shark, herring, and dolphin; while fish such as the mackerel and sergeant fish are both countershaded and patterned with stripes or spots.[20]

It tones the canvas on which are painted the Leopard's spots, the Tiger's stripes ... It is the dress almost universally worn by rodents... It is the essential uniform adopted by Conies, Asses, Antelopes, Deer ... It is repeated extensively among the marsupials ... It provides a basic livery for the great majority of snakes, lizards, and amphibians. Among insects it reaches a fine state of perfection in different caterpillars and grasshoppers. ... It is, however, in rivers, and in the surface waters of the sea, that countershading reaches its maximum development and significance.

— Hugh Cott[18]

Mesozoic marine reptiles had countershading. Fossilised skin pigmented with dark-coloured eumelanin reveals that ichthyosaurs, leatherback turtles and mosasaurs had dark backs and light bellies.[21][22] The ornithischian dinosaur Psittacosaurus similarly appears to have been countershaded, implying that its predators detected their prey by deducing shape from shading. Modelling suggests further that the dinosaur was optimally countershaded for a closed habitat such as a forest.[23]

Counter-illumination

Another form of animal camouflage uses bioluminescence to increase the average brightness of an animal to match the brightness of the background.[24] This is called counter-illumination. It is common in mid-water pelagic fish and invertebrates especially squid. It makes the counter-illuminated animal practically invisible to predators viewing it from below.[25] As such, counter-illumination camouflage can be seen as an extension beyond what countershading can achieve. Where countershading only paints out shadows, counter-illumination can add in actual lights, permitting effective camouflage in changing conditions, including where the background is bright enough to make an animal that is not counter-illuminated appear as a shadow.[26]

Military

Countershading, like counter-illumination, has rarely been applied in practice for military camouflage, though not because military authorities were unaware of it. Both Abbott Thayer in the First World War and Hugh Cott in the Second World War proposed countershading to their countries' armed forces. They each demonstrated the effectiveness of countershading, without succeeding in persuading their armed forces to adopt the technique, though they influenced military adoption of camouflage in general.[11]

Cott was a protege of John Graham Kerr who had quarrelled with Norman Wilkinson in the First World War about dazzle camouflage for ships. Wilkinson remained influential in 1939 as an inspector of camouflage, so a political argument developed. Cott was invited to camouflage a 12-inch rail-mounted gun, alongside a similar gun camouflaged conventionally. Cott carefully combined disruptive contrast to break up the gun barrel's outlines with countershading to flatten out its appearance as a solid cylinder. The guns were then photographed from the air from various angles, and in Peter Forbes's view "the results were remarkable."[27] Cott's gun is "invisible except to the most minute scrutiny by someone who knows exactly where to look and what to look for. The other gun is always highly visible." The authorities hesitated, appearing to be embarrassed by the evidence that Cott was right, and argued that countershading would be too difficult to use as an expert zoologist would be needed to supervise every installation. Cott was posted to the Middle East, and Kerr unsuccessfully intervened, pleading for guns to be painted Cott's way and Cott to be brought home.[28]

The Australian zoologist William Dakin in his 1941 book The Art of Camouflage followed Thayer in describing countershading in some detail, and the book was reprinted as a military handbook in 1942. Dakin photographed model birds, much as Thayer and Cott had done, and argued that the shoulders and arms of battledress should be countershaded.[29]

Countershading was described in the US War Department's 1943 Principles of Camouflage, where after four paragraphs of theory and one on its use in nature, the advice given is that:[30]

Upper surfaces should be painted and textured so as to conform to the color and tone of the surrounding country (background) and the sides graded and toned from this to the white which the under surfaces and parts in shade should be painted.[30]

Inventors have continued to advocate military usage of countershading, with for example a 2005 US patent for personal camouflage including countershading in the form of "statistical countercoloring" with varying sizes of rounded dark patches on a lighter ground.[31]

Research by Ariel Tankus and Yehezkel Yeshurun investigating "camouflage breaking", the automated detection of objects such as tanks, showed that analysing images for convexity by looking for graded shadows can "break very strong camouflage, which might delude even human viewers." More precisely, images are searched for places where the gradient of brightness crosses zero, such as the line where a shadow stops becoming darker and starts to become lighter again. The technique defeated camouflage using disruption of edges, but the authors observed that animals with Thayer countershading are using "counter-measures to convexity based detectors", which implied "predators who use convexity based detectors."[32]

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Ibex

Ibex

An ibex is any of several species of wild goat , distinguished by the male's large recurved horns, which are transversely ridged in front. Ibex are found in Eurasia, North Africa and East Africa. The name ibex comes from Latin, borrowed from Iberian or Aquitanian, akin to Old Spanish bezerro "bull", modern Spanish becerro "yearling". Ranging in height from 70 to 110 centimetres (27–43 in) and weighing 90 to 120 kilograms (200–270 lb), ibex can live up to 20 years. Two closely related varieties of goats found in the wild are not usually called ibex: the markhor and the feral goat.

Deer

Deer

Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including muntjac, elk (wapiti), red deer, and fallow deer; and the Capreolinae, including reindeer (caribou), white-tailed deer, roe deer, and moose. Male deer of all species, as well as female reindeer, grow and shed new antlers each year. In this, they differ from permanently horned antelope, which are part of a different family (Bovidae) within the same order of even-toed ungulates (Artiodactyla).

Blue shark

Blue shark

The blue shark, also known as the great blue shark, is a species of requiem shark, in the family Carcharhinidae, which inhabits deep waters in the world's temperate and tropical oceans. Averaging around 3.1 m (10 ft) and preferring cooler waters, the blue shark migrates long distances, such as from New England to South America. It is listed as Near Threatened by the IUCN.

Herring

Herring

Herring are forage fish, mostly belonging to the family of Clupeidae.

Cobia

Cobia

The cobia is a species of carangiform marine fish, the only extant representative of the genus Rachycentron and the family Rachycentridae. Its other common names include black kingfish, black salmon, ling, lemonfish, crabeater, prodigal son, codfish, and black bonito.

Mesozoic

Mesozoic

The Mesozoic Era is the second-to-last era of Earth's geological history, lasting from about 252 to 66 million years ago, comprising the Triassic, Jurassic and Cretaceous Periods. It is characterized by the dominance of archosaurian reptiles, like the dinosaurs; an abundance of conifers and ferns; a hot greenhouse climate; and the tectonic break-up of Pangaea. The Mesozoic is the middle of the three eras since complex life evolved: the Paleozoic, the Mesozoic, and the Cenozoic.

Fossil

Fossil

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

Ichthyosaur

Ichthyosaur

Ichthyosaurs are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia.

Mosasaur

Mosasaur

Mosasaurs comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on the Meuse in 1764. They belong to the order Squamata, which includes lizards and snakes.

Dinosaur

Dinosaur

Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 245 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is a subject of active research. They became the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201.3 mya and their dominance continued throughout the Jurassic and Cretaceous periods. The fossil record shows that birds are feathered dinosaurs, having evolved from earlier theropods during the Late Jurassic epoch, and are the only dinosaur lineage known to have survived the Cretaceous–Paleogene extinction event approximately 66 mya. Dinosaurs can therefore be divided into avian dinosaurs—birds—and the extinct non-avian dinosaurs, which are all dinosaurs other than birds.

Counter-illumination

Counter-illumination

Counter-illumination is a method of active camouflage seen in marine animals such as firefly squid and midshipman fish, and in military prototypes, producing light to match their backgrounds in both brightness and wavelength.

Bioluminescence

Bioluminescence

Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some bioluminescent bacteria, and terrestrial arthropods such as fireflies. In some animals, the light is bacteriogenic, produced by symbiotic bacteria such as those from the genus Vibrio; in others, it is autogenic, produced by the animals themselves.

Function

Countershading acts as a form of camouflage by 'painting out' the self-shadowing of the body or object. The result is a 'flat' appearance, instead of the 'solid' appearance (with visual convexity) of the body before countershading.
Countershading acts as a form of camouflage by 'painting out' the self-shadowing of the body or object. The result is a 'flat' appearance, instead of the 'solid' appearance (with visual convexity) of the body before countershading.

Hannah Rowland, reviewing countershading 100 years after Abbott Thayer, observed that countershading, which she defines as "darker pigmentation on those surfaces exposed to the most lighting" is a common but poorly understood aspect of animal coloration.[5] She noted there had been "much debate" about how countershading works.[33] She considered the evidence for Thayer's theory that this acts as camouflage "by reducing ventral shadowing", and reviewed alternative explanations for countershading.[5]

Camouflage theories of countershading, Rowland wrote, include "self-shadow concealment which results in improved background matching when viewed from the side"; "self-shadow concealment that flattens the form when viewed from the side"; "background matching when viewed from above or below"; and "body outline obliteration when viewed from above".[5] These are examined in turn below.

Flattening and background matching when viewed from the side

When oriented horizontally, the countershading of the gray squirrel, Sciurus carolinensis, helps to "paint out" its ventral shadow.When oriented vertically, the gray squirrel's pale belly is conspicuous rather than camouflaged.
When oriented horizontally, the countershading of the gray squirrel, Sciurus carolinensis, helps to "paint out" its ventral shadow.
When oriented horizontally, the countershading of the gray squirrel, Sciurus carolinensis, helps to "paint out" its ventral shadow.When oriented vertically, the gray squirrel's pale belly is conspicuous rather than camouflaged.
When oriented vertically, the gray squirrel's pale belly is conspicuous rather than camouflaged.

Cott, like Thayer, argued that countershading would make animals hard to see from the side, as they would "fade into a ghostly elusiveness".[34] Rowland notes that Cott is here reviewing Thayer's theory and "reinforcing the view that a gradation in shading would act to eliminate the effects of ventral shadowing."[5] Kiltie measured the effect of the countershading of the grey squirrel, Sciurus carolinensis, showing that when the squirrel is horizontal the self-shadowing of the belly is partly concealed, but that when the squirrel is vertical (as when climbing a tree trunk) this effect did not occur.[35]

Thayer's original argument, restated by Cott,[34] was that nature did the exact opposite with countershading that an artist did with paint when creating the illusion of solid three-dimensionality, namely counteracting the effect of shade to flatten out form. Shading is a powerful cue used by animals in different phyla to identify the shapes of objects. Research with chicks showed that they preferred to peck at grains with shadows falling below them (as if illuminated from above), so both humans and birds may make use of shading as a depth cue.[5][36]

Background matching from above or below

The mackerel, Scomber scombrus, like many pelagic fish, is dark above, pale below, camouflaging it against the ocean depths and the bright surface.[c]
The mackerel, Scomber scombrus, like many pelagic fish, is dark above, pale below, camouflaging it against the ocean depths and the bright surface.[c]

A completely different function of animal (and military vehicle) coloration is to camouflage the top and bottom surfaces differently, to match their backgrounds below and above respectively. This was noted, for example, by Frank Evers Beddard in 1892:

Among pelagic fish it is common to find the upper surface dark-coloured and the lower surface white, so that the animal is inconspicuous when seen either from above or below.

— Frank Evers Beddard[37]
Top/bottom countershaded[d] Grumman F6F Hellcat
Top/bottom countershaded[d] Grumman F6F Hellcat

Early researchers including Alfred Russel Wallace,[38] Beddard,[39] Cott[40] and Craik[41] argued that in marine animals including pelagic fish such as marlin and mackerel, as well as dolphins, sharks, and penguins the upper and lower surfaces are sharply distinct in tone, with a dark upper surface and often a nearly white lower surface. They suggested that when seen from the top, the darker dorsal surface of the animal would offer camouflage against the darkness of the deep water below. When seen from below, the lighter ventral area would similarly provide the least possible contrast with the sunlit ocean surface above.[5] There is some evidence for this in birds, where birds that catch fish at a medium depth, rather than at the surface or on the seabed, are more often coloured in this way, and the prey of these birds would see only the underside of the bird.[42] Rowland concluded that each possible role for coloration patterns lumped together as "countershading" needs to be evaluated separately, rather than just assuming it functions effectively.[5]

Outline obliteration from above

Rowland (2009) identified an additional mechanism of countershading not previously analysed, namely that a round body such as a cylinder illuminated and seen from above appears to have dark sides. Using a graphics tool, she demonstrated that this effect can be flattened out by countershading. Since predators are known to use edges to identify prey, countershading may therefore, she argues, make prey harder to detect when seen from above.[5]

Non-camouflage theories

Non-camouflage theories include protection from ultraviolet light; thermoregulation; and protection from abrasion. All three of these "plausible" theories remained largely untested in 2009, according to Rowland.[5]

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Camouflage

Camouflage

Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the battledress of a modern soldier, and the leaf-mimic katydid's wings. A third approach, motion dazzle, confuses the observer with a conspicuous pattern, making the object visible but momentarily harder to locate, as well as making general aiming easier. The majority of camouflage methods aim for crypsis, often through a general resemblance to the background, high contrast disruptive coloration, eliminating shadow, and countershading. In the open ocean, where there is no background, the principal methods of camouflage are transparency, silvering, and countershading, while the ability to produce light is among other things used for counter-illumination on the undersides of cephalopods such as squid. Some animals, such as chameleons and octopuses, are capable of actively changing their skin pattern and colours, whether for camouflage or for signalling. It is possible that some plants use camouflage to evade being eaten by herbivores.

List of camouflage methods

List of camouflage methods

Camouflage is the concealment of animals or objects of military interest by any combination of methods that helps them to remain unnoticed. This includes the use of high-contrast disruptive patterns as used on military uniforms, but anything that delays recognition can be used as camouflage. Camouflage involves deception, whether by looking like the background or by resembling something else, which may be plainly visible to observers. This article lists methods used by animals and the military to escape notice.

Animal coloration

Animal coloration

Animal coloration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male has strong patterns, conspicuous colours and is iridescent, while the female is far less visible.

Gray squirrel

Gray squirrel

Gray squirrel or grey squirrel may refer to several species of squirrel indigenous to North America:The eastern gray squirrel, from the eastern United States and southeastern Canada; introduced into the United Kingdom, Ireland, western North America, Italy, and South Africa The western gray squirrel, from the western United States The Arizona gray squirrel, from the southwestern United States and adjacent Mexico The Mexican gray squirrel, from southern Mexico and Guatemala; introduced into the Florida Keys

Eastern gray squirrel

Eastern gray squirrel

The eastern gray squirrel, also known, particularly outside of North America, as simply the grey squirrel, is a tree squirrel in the genus Sciurus. It is native to eastern North America, where it is the most prodigious and ecologically essential natural forest regenerator. Widely introduced to certain places around the world, the eastern gray squirrel in Europe, in particular, is regarded as an invasive species.

Phylum

Phylum

In biology, a phylum is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia contains about 31 phyla, the plant kingdom Plantae contains about 14 phyla, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships between phyla, which are contained in larger clades, like Ecdysozoa and Embryophyta.

Frank Evers Beddard

Frank Evers Beddard

Frank Evers Beddard FRS FRSE was an English zoologist. He became a leading authority on annelids, including earthworms. He won the Linnean Medal in 1916 for his book on oligochaetes.

Grumman F6F Hellcat

Grumman F6F Hellcat

The Grumman F6F Hellcat is an American carrier-based fighter aircraft of World War II. Designed to replace the earlier F4F Wildcat and to counter the Japanese Mitsubishi A6M Zero, it was the United States Navy's dominant fighter in the second half of the Pacific War. In gaining that role, it prevailed over its faster competitor, the Vought F4U Corsair, which initially had problems with visibility and carrier landings.

Alfred Russel Wallace

Alfred Russel Wallace

Alfred Russel Wallace was an English naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution through natural selection. His 1858 paper on the subject was published that year alongside extracts from Charles Darwin's earlier writings on the topic. It spurred Darwin to set aside the "big species book" he was drafting, and quickly write an abstract of it, published in 1859 as On the Origin of Species.

Marlin

Marlin

Marlins are fish from the family Istiophoridae, which includes about 10 species. A marlin has an elongated body, a spear-like snout or bill, and a long, rigid dorsal fin which extends forward to form a crest. Its common name is thought to derive from its resemblance to a sailor's marlinspike. Marlins are among the fastest marine swimmers. However, greatly exaggerated speeds are often claimed in popular literature, based on unreliable or outdated reports.

Mackerel

Mackerel

Mackerel is a common name applied to a number of different species of pelagic fish, mostly from the family Scombridae. They are found in both temperate and tropical seas, mostly living along the coast or offshore in the oceanic environment.

Dolphin

Dolphin

A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae, Platanistidae, Iniidae, Pontoporiidae, and the extinct Lipotidae. There are 40 extant species named as dolphins.

Evidence

Despite demonstrations and examples adduced by Cott and others, little experimental evidence for the effectiveness of countershading was gathered in the century since Thayer's discovery. Experiments in 2009 using artificial prey showed that countershaded objects do have survival benefits[43] and in 2012, a study by William Allen and colleagues showed that countershading in 114 species of ruminants closely matched predictions for "self-shadow concealment", the function predicted by Poulton, Thayer and Cott.[44]

Mechanism

Evolutionary developmental biology has assembled evidence from embryology and genetics to show how evolution has acted at all scales from the whole organism down to individual genes, proteins and genetic switches. In the case of countershaded mammals with dark (often brownish) upper parts and lighter (often buff or whitish) under parts, such as in the house mouse, it is the Agouti gene which creates the difference in shading. Agouti encodes for a protein, the Agouti signalling peptide (ASP), which specifically inhibits the action of the Melanocortin 1 receptor (MC1R). In the absence of the Agouti protein, alpha-melanocyte-stimulating hormone stimulates the cells bearing MC1R, melanocytes, to produce dark eumelanin, colouring the skin and fur dark brown or black. In the presence of the Agouti protein, the same system produces the lighter-coloured, yellow or red phaeomelanin. A genetic switch active in the cells of the embryo that will become the belly skin causes the Agouti gene to become active there, creating the countershading seen in adult mammals.[45]

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Evolutionary developmental biology

Evolutionary developmental biology

Evolutionary developmental biology is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved.

Embryology

Embryology

Embryology is the branch of animal biology that studies the prenatal development of gametes, fertilization, and development of embryos and fetuses. Additionally, embryology encompasses the study of congenital disorders that occur before birth, known as teratology.

Genetics

Genetics

Genetics is the study of genes, genetic variation, and heredity in organisms. It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar working in the 19th century in Brno, was the first to study genetics scientifically. Mendel studied "trait inheritance", patterns in the way traits are handed down from parents to offspring over time. He observed that organisms inherit traits by way of discrete "units of inheritance". This term, still used today, is a somewhat ambiguous definition of what is referred to as a gene.

Evolution

Evolution

In biology, evolution is the change in heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation tends to exist within any given population as a result of genetic mutation and recombination. Evolution occurs when evolutionary processes such as natural selection and genetic drift act on this variation, resulting in certain characteristics becoming more common or more rare within a population. The evolutionary pressures that determine whether a characteristic is common or rare within a population constantly change, resulting in a change in heritable characteristics arising over successive generations. It is this process of evolution that has given rise to biodiversity at every level of biological organisation.

Gene

Gene

In biology, the word gene can have several different meanings. The Mendelian gene is a basic unit of heredity and the molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protein-coding genes and noncoding genes.

Protein

Protein

Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, responding to stimuli, providing structure to cells and organisms, and transporting molecules from one location to another. Proteins differ from one another primarily in their sequence of amino acids, which is dictated by the nucleotide sequence of their genes, and which usually results in protein folding into a specific 3D structure that determines its activity.

House mouse

House mouse

The house mouse is a small mammal of the order Rodentia, characteristically having a pointed snout, large rounded ears, and a long and almost hairless tail. It is one of the most abundant species of the genus Mus. Although a wild animal, the house mouse has benefited significantly from associating with human habitation to the point that truly wild populations are significantly less common than the semi-tame populations near human activity.

Melanocortin 1 receptor

Melanocortin 1 receptor

The melanocortin 1 receptor (MC1R), also known as melanocyte-stimulating hormone receptor (MSHR), melanin-activating peptide receptor, or melanotropin receptor, is a G protein–coupled receptor that binds to a class of pituitary peptide hormones known as the melanocortins, which include adrenocorticotropic hormone (ACTH) and the different forms of melanocyte-stimulating hormone (MSH). It is coupled to Gαs and upregulates levels of cAMP by activating adenylyl cyclase in cells expressing this receptor. It is normally expressed in skin and melanocytes, and to a lesser degree in periaqueductal gray matter, astrocytes and leukocytes. In skin cancer, MC1R is highly expressed in melanomas but not carcinomas.

Melanocyte-stimulating hormone

Melanocyte-stimulating hormone

The melanocyte-stimulating hormones, known collectively as MSH, also known as melanotropins or intermedins, are a family of peptide hormones and neuropeptides consisting of α-melanocyte-stimulating hormone (α-MSH), β-melanocyte-stimulating hormone (β-MSH), and γ-melanocyte-stimulating hormone (γ-MSH) that are produced by cells in the pars intermedia of the anterior lobe of the pituitary gland.

Melanocyte

Melanocyte

Melanocytes are melanin-producing neural crest-derived cells located in the bottom layer of the skin's epidermis, the middle layer of the eye, the inner ear, vaginal epithelium, meninges, bones, and heart. Melanin is a dark pigment primarily responsible for skin color. Once synthesized, melanin is contained in special organelles called melanosomes which can be transported to nearby keratinocytes to induce pigmentation. Thus darker skin tones have more melanosomes present than lighter skin tones. Functionally, melanin serves as protection against UV radiation. Melanocytes also have a role in the immune system.

Embryo

Embryo

An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm cell. The resulting fusion of these two cells produces a single-celled zygote that undergoes many cell divisions that produce cells known as blastomeres. The blastomeres are arranged as a solid ball that when reaching a certain size, called a morula, takes in fluid to create a cavity called a blastocoel. The structure is then termed a blastula, or a blastocyst in mammals.

Reverse countershading

The honey badger is reverse countershaded, a form of aposematism (warning coloration).
The honey badger is reverse countershaded, a form of aposematism (warning coloration).
The sea slug Glaucus atlanticus swims and is countershaded upside-down.
The sea slug Glaucus atlanticus swims and is countershaded upside-down.

If countershading paints out shadows, the reverse, darkening the belly and lightening the back, would maximise contrast by adding to the natural fall of light. This pattern of animal coloration is found in animals such as the skunk and honey badger with strong defences—the offensive stink of the skunk, and the sharp claws, aggressive nature and stink of the honey badger.[46] These animals do not run when under attack, but move slowly, often turning to face the danger, and giving deimatic or threat displays either to startle inexperienced predators, or as an aposematic signal, to warn off experienced ones.[47]

The caterpillar of the Luna moth, as discovered by Thayer, is in Cott's phrase "countershaded in relation to [its] attitude", i.e. shaded with a light back grading to a dark belly, as is the Nile catfish, Synodontis batensoda for the same reason: these animals (and other caterpillars including Automeris io and the eyed hawkmoth, Smerinthus ocellatus) habitually live 'upside down' with the belly uppermost. Similarly in the sea slug Glaucus atlanticus, the reverse countershading is associated with inverted habits. These animals are thus employing countershading in the usual way for camouflage.[48]

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Honey badger

Honey badger

The honey badger, also known as the ratel, is a mammal widely distributed in Africa, Southwest Asia, and the Indian subcontinent. Because of its wide range and occurrence in a variety of habitats, it is listed as Least Concern on the IUCN Red List.

Aposematism

Aposematism

Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or smell, sharp spines, or aggressive nature. These advertising signals may take the form of conspicuous coloration, sounds, odours, or other perceivable characteristics. Aposematic signals are beneficial for both predator and prey, since both avoid potential harm.

Glaucus atlanticus

Glaucus atlanticus

Glaucus atlanticus is a species of small, blue sea slug, a pelagic (open-ocean) aeolid nudibranch, a shell-less gastropod mollusk in the family Glaucidae.

Animal coloration

Animal coloration

Animal coloration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male has strong patterns, conspicuous colours and is iridescent, while the female is far less visible.

Skunk

Skunk

Skunks are mammals in the family Mephitidae. They are known for their ability to spray a liquid with a strong, unpleasant scent from their anal glands. Different species of skunk vary in appearance from black-and-white to brown, cream or ginger colored, but all have warning coloration.

Deimatic behaviour

Deimatic behaviour

Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey animal an opportunity to escape. The term deimatic or dymantic originates from the Greek δειματόω (deimatóo), meaning "to frighten".

Synodontis batensoda

Synodontis batensoda

Synodontis batensoda, the upside-down catfish, is a species of mochokid upside-down catfish. It is unevenly distributed in inland waters across Africa from Senegal to Ethiopia, and is also known as a squeaker or giant upside-down catfish. It was originally described by Eduard Rüppell in 1832 in the paper "Continuation of the description and figure of several new fish, in the Nile. p1-14".

Automeris io

Automeris io

Automeris io, the Io moth or peacock moth, is a colorful North American moth in the family Saturniidae. The io moth is also a member of the subfamily Hemileucinae. The name Io comes from Greek mythology in which Io was a mortal lover of Zeus. The Io moth ranges from the southeast corner of Manitoba and in the southern extremes of Ontario, Quebec, New Brunswick and Nova Scotia in Canada, and in the US it is found from Montana, North Dakota, South Dakota, Nebraska, Colorado, New Mexico, Texas, Utah, east of those states and down to the southern end of Florida. The species was first described by Johan Christian Fabricius in 1775.

Smerinthus ocellatus

Smerinthus ocellatus

Smerinthus ocellatus, the eyed hawk-moth, is a European moth of the family Sphingidae. The species was first described by Carl Linnaeus in his 1758 10th edition of Systema Naturae.

Examples in animals

Source: "Countershading", Wikipedia, Wikimedia Foundation, (2023, February 20th), https://en.wikipedia.org/wiki/Countershading.

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See also
Notes
  1. ^ It was called Rumia crataegata at the time.
  2. ^ It was called Amphidasis betularia at the time.
  3. ^ The mackerel, like many other pelagic fish, is also camouflaged by silvering, and when seen from above it has a bold disruptive pattern.
  4. ^ Colours used are Non-Specular Sea Blue, Intermediate Blue, White.
References
  1. ^ Argo, Emily (21 April 2017). "Countershading". Fishionary. American Fisheries Society. Retrieved 17 December 2022.
  2. ^ Poulton, 1888.
  3. ^ Poulton, 1887.
  4. ^ Thayer, 1909. p 22.
  5. ^ a b c d e f g h i j Rowland, 2009.
  6. ^ Forbes, 2009. pp. 76–79.
  7. ^ Thayer, 1909.
  8. ^ Thayer, 1909. pp 14–15.
  9. ^ Thayer, 1909. Plate XII.
  10. ^ U.S. Patent 715,013
  11. ^ a b Goldstein, 2009, pp. 233–235.
  12. ^ Cott, 1940. pp. 35–46.
  13. ^ Cott, 1940. pp. 172–173.
  14. ^ Thayer, 1909. Caption to Figure 7.
  15. ^ Cott, 1940. Caption to Plate 7.
  16. ^ Cott, 1940. p. 35.
  17. ^ ONR, 2013.
  18. ^ a b c Cott, 1940. p. 40.
  19. ^ Cott, 1940. p37.
  20. ^ Cott, 1940. p41
  21. ^ Lindgren, Johan and Peter Sjövall, Ryan M. Carney, Per Uvdal, Johan A. Gren, Gareth Dyke, Bo Pagh Schultz, Matthew D. Shawkey, Kenneth R. Barnes, Michael J. Polcyn (February 2014). "Skin pigmentation provides evidence of convergent melanism in extinct marine reptiles". Nature. 506 (7489): 484–488. Bibcode:2014Natur.506..484L. doi:10.1038/nature12899. PMID 24402224. S2CID 4468035.{{cite journal}}: CS1 maint: uses authors parameter (link)
  22. ^ Lindgren, Johan; Sjövall, Peter; Thiel, Volker; Zheng, Wenxia; Ito, Shosuke; Wakamatsu, Kazumasa; Hauff, Rolf; Kear, Benjamin P.; Engdahl, Anders; Alwmark, Carl; Eriksson, Mats E.; Jarenmark, Martin; Sachs, Sven; Ahlberg, Per E.; Marone, Federica (December 2018). "Soft-tissue evidence for homeothermy and crypsis in a Jurassic ichthyosaur". Nature. 564 (7736): 359–365. Bibcode:2018Natur.564..359L. doi:10.1038/s41586-018-0775-x. ISSN 1476-4687. PMID 30518862. S2CID 54458324.
  23. ^ Vinther, Jakob; Nicholls, Robert; Lautenschlager, Stephan; Pittman, Michael; Kaye, Thomas G.; Rayfield, Emily; Mayr, Gerald; Cuthill, Innes C. (2016). "3D Camouflage in an Ornithischian Dinosaur". Current Biology. 26 (18): 2456–2462. doi:10.1016/j.cub.2016.06.065. PMC 5049543. PMID 27641767.
  24. ^ Young and Roper, 1977.
  25. ^ Young and Roper, 1976.
  26. ^ Jones, 2004. p. 1151.
  27. ^ Forbes, 2009. p. 150.
  28. ^ a b Forbes, 2009. pp. 142–146, 149–151, 156.
  29. ^ Elias, 2011.
  30. ^ a b Anon, 1943.
  31. ^ Tooley, 2005.
  32. ^ Tankus and Yeshurun, 2013.
  33. ^ Rowland, 2011.
  34. ^ a b Cott, 1940. pp 36–37.
  35. ^ Kiltie, 1944.
  36. ^ Hershberger, 1970.
  37. ^ Beddard, 1892. p. 122.
  38. ^ Wallace, 1889, p 193
  39. ^ Beddard, 1895, p 115
  40. ^ Cott, 1940.
  41. ^ Craik, 1944.
  42. ^ Ruxton 2004.
  43. ^ Rowland et al, 2009.
  44. ^ Allen et al, 2012.
  45. ^ Carroll, Sean B. (2006). Endless Forms Most Beautiful. Weidenfeld and Nicolson. pp. 229–231, 237. ISBN 978-0-297-85094-6.
  46. ^ "Black, White and Stinky: Explaining Coloration in Skunks and Other Boldly Colored Animals". University of Massachusetts Amherst. 27 May 2011. Archived from the original on 15 October 2014. Retrieved 19 June 2014.
  47. ^ Edmunds, 2008.
  48. ^ Cott, 1940. p. 43.
Bibliography

Pioneering books

General reading

  • Behrens, Roy R. (2009). Goldstein, E Bruce (ed.). Encyclopedia of Perception, Volume 1. Sage. pp. 233–235.
  • Edmunds, Malcolm (2008). "Deimatic Behavior". In Capinera, John L. (ed.). Encyclopedia of Entomology. Springer. ISBN 9781402062421.
  • Forbes, Peter (2009). Dazzled and Deceived: Mimicry and Camouflage. Yale.
  • Rowland, Hannah M. (2011). "The history, theory and evidence for a cryptic function of countershading". In Stevens, Martin; Merilaita, Sami (eds.). Animal Camouflage: Mechanisms and Function. Cambridge University Press.
  • Ruxton, Graeme D.; Sherratt, Thomas N.; Speed, Michael P. (2004). "3. Countershading and counterillumination". Avoiding Attack: The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry. Oxford University Press.

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