Thepeacock flounder can change its pattern and colours to match its environment.A soldier applying camouflage face paint; both helmet and jacket aredisruptively patterned.
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 theleopard's spotted coat, thebattledress of a modernsoldier, and theleaf-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. The majority of camouflage methods aim for crypsis, often through a general resemblance to the background, high contrastdisruptive coloration, eliminating shadow, andcountershading. In the open ocean, where there is no background, the principal methods of camouflage are transparency, silvering, and countershading, while theability to produce light is among other things used forcounter-illumination on the undersides ofcephalopods such assquid. Some animals, such aschameleons andoctopuses, are capable ofactively changing their skin pattern and colours, whether for camouflage or for signalling. It is possible that some plants use camouflage to evade being eaten byherbivores.
Military camouflage was spurred by the increasing range and accuracy of firearms in the 19th century. In particular the replacement of the inaccuratemusket with therifle made personal concealment in battle a survival skill. In the 20th century, military camouflage developed rapidly, especially during theWorld War I. On land, artists such asAndré Mare designed camouflage schemes and observation posts disguised as trees.At sea, merchant ships and troop carriers were painted indazzle patterns that were highly visible, but designed to confuse enemy submarines as to the target's speed, range, and heading. During and afterWorld War II, a variety of camouflage schemes were used foraircraft and for ground vehicles in different theatres of war. The use of radar since the mid-20th century has largely made camouflage for fixed-wing military aircraft obsolete.
Non-military use of camouflage includes makingcell telephone towers less obtrusive and helping hunters to approach wary game animals. Patterns derived from military camouflage are frequently used in fashion clothing, exploiting their strong designs and sometimes their symbolism. Camouflage themes recur in modern art, and both figuratively and literally in science fiction and works of literature.
Camouflage has been a topic of interest and research inzoology for well over a century. According toCharles Darwin's 1859 theory ofnatural selection,[2] features such as camouflageevolved by providing individual animals with a reproductive advantage, enabling them to leave more offspring, on average, than other members of the samespecies. In hisOrigin of Species, Darwin wrote:[3]
When we see leaf-eating insects green, and bark-feeders mottled-grey; the alpine ptarmigan white in winter, thered-grouse the colour ofheather, and theblack-grouse that ofpeaty earth, we must believe that these tints are of service to these birds and insects in preserving them from danger. Grouse, if not destroyed at some period of their lives, would increase in countless numbers; they are known to suffer largely frombirds of prey; and hawks are guided by eyesight to their prey, so much so, that on parts of the Continent persons are warned not to keep white pigeons, as being the most liable to destruction. Hence I can see no reason to doubt that natural selection might be most effective in giving the proper colour to each kind of grouse, and in keeping that colour, when once acquired, true and constant.[3]
The English zoologistEdward Bagnall Poulton studiedanimal coloration, especially camouflage. In his 1890 bookThe Colours of Animals, he classified different types such as "special protective resemblance" (where an animal looks like another object), or "general aggressive resemblance" (where a predator blends in with the background, enabling it to approach prey). His experiments showed thatswallow-tailed mothpupae were camouflaged to match the backgrounds on which they were reared aslarvae.[4][a] Poulton's "general protective resemblance"[6] was at that time considered to be the main method of camouflage, as whenFrank Evers Beddard wrote in 1892 that "tree-frequenting animals are often green in colour. Among vertebrates numerous species ofparrots,iguanas,tree-frogs, and thegreen tree-snake are examples".[7] Beddard did however briefly mention other methods, including the "alluring coloration" of theflower mantis and the possibility of a different mechanism in theorange tip butterfly. He wrote that "the scattered green spots upon the under surface of the wings might have been intended for a rough sketch of the small flowerets of the plant [anumbellifer], so close is their mutual resemblance."[8][b] He also explained the coloration of sea fish such as themackerel: "Amongpelagic 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."[10]
Abbott Thayer's 1907 paintingPeacock in the Woods depicted apeacock as if it were camouflaged.
The artistAbbott Handerson Thayer formulated what is sometimes called Thayer's Law, the principle ofcountershading.[11] However, he overstated the case in the 1909 bookConcealing-Coloration in the Animal Kingdom, arguing that "All patterns and colors whatsoever of all animals that ever preyed or are preyed on are under certain normal circumstances obliterative" (that is, cryptic camouflage), and that "Not one 'mimicry' mark, not one 'warning color'... nor any 'sexually selected' color, exists anywhere in the world where there is not every reason to believe it the very best conceivable device for the concealment of its wearer",[12][13] and using paintings such asPeacock in the Woods (1907) to reinforce his argument.[14] Thayer was roundly mocked for these views by critics includingTeddy Roosevelt.[15]
The English zoologistHugh Cott's 1940 bookAdaptive Coloration in Animals corrected Thayer's errors, sometimes sharply: "Thus we find Thayer straining the theory to a fantastic extreme in an endeavour to make it cover almost every type of coloration in the animal kingdom."[16] Cott built on Thayer's discoveries, developing a comprehensive view of camouflage based on "maximum disruptive contrast", countershading and hundreds of examples. The book explained howdisruptive camouflage worked, using streaks of boldly contrasting colour, paradoxically making objects less visible by breaking up their outlines.[17] While Cott was more systematic and balanced in his view than Thayer, and did include some experimental evidence on the effectiveness of camouflage,[18] his 500-page textbook was, like Thayer's, mainly anatural history narrative which illustrated theories with examples.[19]
Experimental evidence that camouflage helps prey avoid being detected bypredators was first provided in 2016, when ground-nesting birds (plovers andcoursers) were shown to survive according to how well their egg contrast matched the local environment.[20]
As there is a lack of evidence for camouflage in the fossil record, studying the evolution of camouflage strategies is very difficult. Furthermore, camouflage traits must be both adaptable (provide a fitness gain in a given environment) and heritable (in other words, the trait must undergopositive selection).[21] Thus, studying the evolution of camouflage strategies requires an understanding of the genetic components and various ecological pressures that drive crypsis.
Camouflage is a soft-tissue feature that is rarely preserved in thefossil record, but rare fossilised skin samples from theCretaceous period show that some marine reptiles were countershaded. The skins, pigmented with dark-colouredeumelanin, reveal that bothleatherback turtles andmosasaurs had dark backs and light bellies.[22] There is fossil evidence of camouflaged insects going back over 100 million years, for example lacewings larvae that stick debris all over their bodies much as their modern descendants do, hiding them from their prey.[23] Dinosaurs appear to have been camouflaged, as a 120 million year old fossil of aPsittacosaurus has been preserved withcountershading.[24]
Camouflage does not have a single genetic origin. However, studying the genetic components of camouflage in specific organisms illuminates the various ways that crypsis can evolve among lineages. Manycephalopods have the ability to actively camouflage themselves, controlling crypsis through neural activity. For example, the genome of the common cuttlefish includes 16 copies of thereflectin gene, which grants the organism remarkable control over coloration and iridescence.[25] The reflectin gene is thought to have originated through transposition from symbioticAliivibrio fischeri bacteria, which provide bioluminescence to its hosts. While not all cephalopods useactive camouflage, ancient cephalopods may have inherited the gene horizontally from symbioticA. fischeri, with divergence occurred through subsequent gene duplication (such as in the case ofSepia officinalis) or gene loss (as with cephalopods with no active camouflage capabilities).[26][3] This is unique as an instance of camouflage arising as an instance ofhorizontal gene transfer from anendosymbiont. However, other methods of horizontal gene transfer are common in the evolution of camouflage strategies in other lineages.Peppered moths andwalking stick insects both have camouflage-related genes that stem from transposition events.[27][28]
TheAgouti genes are orthologous genes involved in camouflage across many lineages. They produce yellow and red coloration (phaeomelanin), and work in competition with other genes that produce black (melanin) and brown (eumelanin) colours.[29] Ineastern deer mice, over a period of about 8000 years the single agouti gene developed 9 mutations that each made expression of yellow fur stronger under natural selection, and largely eliminated melanin-coding black fur coloration.[30] On the other hand, all blackdomesticated cats have deletions of the agouti gene that prevent its expression, meaning no yellow or red color is produced. The evolution, history and widespread scope of the agouti gene shows that different organisms often rely on orthologous or even identical genes to develop a variety of camouflage strategies.[31]
While camouflage can increase an organism's fitness, it has genetic and energetic costs. There is a trade-off between detectability and mobility. Species camouflaged to fit a specificmicrohabitat are less likely to be detected when in that microhabitat, but must spend energy to reach, and sometimes to remain in, such areas. Outside the microhabitat, the organism has a higher chance of detection. Generalized camouflage allows species to avoid predation over a wide range of habitat backgrounds, but is less effective. The development of generalized or specialized camouflage strategies is highly dependent on the biotic and abiotic composition of the surrounding environment.[32]
There are many examples of the tradeoffs between specific and general cryptic patterning.Phestilla melanocrachia, a species of nudibranch that feeds onstony coral, utilizes specific cryptic patterning in reef ecosystems. The nudibranch syphons pigments from the consumed coral into the epidermis, adopting the same shade as the consumed coral. This allows the nudibranch to change colour (mostly between black and orange) depending on the coral system that it inhabits. However,P. melanocrachia can only feed and lay eggs on the branches of host-coral,Platygyra carnosa, which limits the geographical range and efficacy in nudibranch nutritional crypsis. Furthermore, the nudibranch colour change is not immediate, and switching between coral hosts when in search for new food or shelter can be costly.[33]
The costs associated with distractive or disruptive crypsis are more complex than the costs associated with background matching. Disruptive patterns distort the body outline, making it harder to precisely identify and locate.[34] However, disruptive patterns result in higher predation.[35] Disruptive patterns that specifically involve visible symmetry (such as in some butterflies) reduce survivability and increase predation.[36] Some researchers argue that because wing-shape and color pattern are genetically linked, it is genetically costly to develop asymmetric wing colorations that would enhance the efficacy of disruptive cryptic patterning. Symmetry does not carry a high survival cost for butterflies and moths that their predators views from above on a homogeneous background, such as the bark of a tree. On the other hand, natural selection drives species with variable backgrounds and habitats to move symmetrical patterns away from the centre of the wing and body, disrupting their predators' symmetry recognition.[37]
Camouflage can be achieved by different methods, described below. Most of the methods help to hide against a background; but mimesis and motion dazzle protect without hiding. Methods may be applied on their own or in combination. Many mechanisms are visual, but some research has explored the use of techniques againstolfactory (scent) andacoustic (sound) detection.[38][39] Methods may also apply to military equipment.[40]
Some animals' colours and patterns match a particular natural background. This is an important component of camouflage in all environments. For instance, tree-dwellingparakeets are mainly green;woodcocks of the forest floor are brown and speckled; reedbedbitterns are streaked brown and buff; in each case the animal's coloration matches the hues of its habitat.[41][42] Similarly,desert animals are almost all desert coloured in tones of sand, buff, ochre, and brownish grey, whether they are mammals like thegerbil orfennec fox, birds such as thedesert lark orsandgrouse, or reptiles like theskink orhorned viper.[43] Military uniforms, too, generally resemble their backgrounds; for examplekhaki uniforms are a muddy or dusty colour, originally chosen for service in South Asia.[44] Many moths showindustrial melanism,[45] including thepeppered moth which has coloration that blends in with tree bark.[46] The coloration of these insectsevolved between 1860 and 1940 to match the changing colour of the tree trunks on which they rest, from pale and mottled to almost black in polluted areas.[45][c] This is taken by zoologists asevidence that camouflage is influenced by natural selection, as well as demonstrating that it changes where necessary to resemble the local background.[45]
Lion in Kruger National Park, South Africa, blending in with the tall grass
Illustration of the principle of "maximum disruptive contrast" byHugh Cott, 1940
Disruptive patterns use strongly contrasting, non-repeating markings such as spots or stripes to break up the outlines of an animal or military vehicle,[47] or to conceal telltale features, especiallyby masking the eyes, as in thecommon frog.[48] Disruptive patterns may use more than one method to defeat visual systems such asedge detection.[49] Predators like theleopard use disruptive camouflage to help them approach prey, while potential prey use it to avoid detection by predators.[50] Disruptive patterning is common in military usage, both for uniforms and for military vehicles. Disruptive patterning, however, does not always achieve crypsis on its own, as an animal or a military target may be given away by factors like shape, shine, and shadow.[51][52][53]
The presence of bold skin markings does not in itself prove that an animal relies on camouflage, as that depends on its behaviour.[54] For example, althoughgiraffes have a high contrast pattern that could be disruptive coloration, the adults are very conspicuous when in the open. Some authors have argued that adult giraffes are cryptic, since when standing among trees and bushes they are hard to see at even a few metres' distance.[55] However, adult giraffes move about to gain the best view of an approaching predator, relying on their size and ability to defend themselves, even from lions, rather than on camouflage.[55] A different explanation is implied by young giraffes being far more vulnerable to predation than adults. More than half of all giraffe calves die within a year,[55] and giraffe mothers hide their newly born calves, which spend much of the time lying down in cover while their mothers are away feeding. The mothers return once a day to feed their calves with milk. Since the presence of a mother nearby does not affect survival, it is argued that these juvenile giraffes must be very well camouflaged; this is supported by coat markings being stronglyinherited.[55]
The possibility ofcamouflage in plants was little studied until the late 20th century. Leafvariegation with white spots may serve as camouflage in forestunderstory plants, where there is a dappled background; leaf mottling is correlated with closed habitats. Disruptive camouflage would have a clear evolutionary advantage in plants: they would tend to escape from being eaten byherbivores. Another possibility is that some plants have leaves differently coloured on upper and lower surfaces or on parts such as veins and stalks to make green-camouflaged insects conspicuous, and thus benefit the plants by favouring the removal of herbivores by carnivores. These hypotheses are testable.[56][57][58]
Leopard: a disruptively camouflaged predator
Russian T-90 battle tank painted in bold disruptive pattern of sand and green
Gaboon viper's bold markings are powerfully disruptive.
Aptarmigan and five chicks exhibit exceptional disruptive camouflage
Jumping spider: a disruptively camouflaged invertebrate predator
Manyunderstory plants such as the saw greenbriar,Smilax bona-nox have pale markings, possibly disruptive camouflage.
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 of the body before countershading.
Countershading uses graded colour to counteract the effect of self-shadowing, creating an illusion of flatness. Self-shadowing makes an animal appear darker below than on top, grading from light to dark; countershading 'paints in' tones which are darkest on top, lightest below, making the countershaded animal nearly invisible against a suitable background.[59] Thayer observed that "Animals are painted by Nature, darkest on those parts which tend to be most lighted by the sky's light, andvice versa". Accordingly, the principle of countershading is sometimes calledThayer's Law.[60] Countershading is widely used byterrestrial animals, such asgazelles[61] and grasshoppers; marine animals, such assharks anddolphins;[62] and birds, such assnipe anddunlin.[63][64]
Countershading is less often used for military camouflage, despite Second World War experiments that showed its effectiveness. EnglishzoologistHugh Cott encouraged the use of methods including countershading, but despite his authority on the subject, failed to persuade the British authorities.[65] Soldiers often wrongly viewed camouflage netting as a kind of invisibility cloak, and they had to be taught to look at camouflage practically, from an enemy observer's viewpoint.[66][67] At the same time inAustralia, zoologistWilliam John Dakin advised soldiers to copy animals' methods, using their instincts for wartime camouflage.[68]
The term countershading has a second meaning unrelated to "Thayer's Law". It is that the upper and undersides of animals such as sharks, and of some military aircraft, are different colours to match the different backgrounds when seen from above or from below. Here the camouflage consists of two surfaces, each with the simple function of providing concealment against a specific background, such as a bright water surface or the sky. The body of a shark or the fuselage of an aircraft is not gradated from light to dark to appear flat when seen from the side. The camouflage methods used are the matching of background colour and pattern, and disruption of outlines.[61]
Camouflaged animals and vehicles are readily given away by their shapes and shadows. A flange helps to hide the shadow and a pale fringe breaks up and averages out any shadow that remains.
Some animals, such as thehorned lizards of North America, have evolved elaborate measures to eliminateshadow. Their bodies are flattened, with the sides thinning to an edge; the animals habitually press their bodies to the ground; and their sides are fringed with white scales which effectively hide and disrupt any remaining areas of shadow there may be under the edge of the body.[69] The theory that the body shape of the horned lizards which live in open desert is adapted to minimise shadow is supported by the one species which lacks fringe scales, theroundtail horned lizard, which lives in rocky areas and resembles a rock. When this species is threatened, it makes itself look as much like a rock as possible by curving its back, emphasizing its three-dimensional shape.[69] Some species of butterflies, such as the speckled wood,Pararge aegeria, minimise their shadows when perched by closing the wings over their backs, aligning their bodies with the sun, and tilting to one side towards the sun, so that the shadow becomes a thin inconspicuous line rather than a broad patch.[70] Similarly, some ground-nesting birds, including theEuropean nightjar, select a resting position facing the sun.[70] Eliminating shadow was identified as a principle of military camouflage during theSecond World War.[71]
Three countershaded and cryptically colouredibex almost invisible in the Israeli desert
"Shape, shine, shadow" make these 'camouflaged' military vehicles easily visible.
Many prey animals have conspicuous high-contrast markings which paradoxically attract the predator's gaze.[d][72] Thesedistractive markings may serve as camouflage by distracting the predator's attention from recognising the prey as a whole, for example by keeping the predator from identifying the prey's outline. Experimentally, search times forblue tits increased when artificial prey had distractive markings.[73]
Theleafy sea dragon sways like seaweeds to reinforce its camouflage.
Movement catches the eye of prey animals on the lookout for predators, and of predators hunting for prey.[74] Most methods of crypsis therefore also require suitable cryptic behaviour, such as lying down and keeping still to avoid being detected, or in the case of stalking predators such as thetiger, moving with extreme stealth, both slowly and quietly, watching its prey for any sign they are aware of its presence.[74] As an example of the combination of behaviours and other methods of crypsis involved, young giraffes seek cover, lie down, and keep still, often for hours until their mothers return; their skin pattern blends with the pattern of the vegetation, while the chosen cover and lying position together hide the animals' shadows.[55] Theflat-tail horned lizard similarly relies on a combination of methods: it is adapted to lie flat in the open desert, relying on stillness, its cryptic coloration, and concealment of its shadow to avoid being noticed by predators.[75] In the ocean, theleafy sea dragon sways mimetically, like the seaweeds amongst which it rests, as if rippled by wind or water currents.[76] Swaying is seen also in some insects, like Macleay's spectre stick insect,Extatosoma tiaratum. The behaviour may be motion crypsis, preventing detection, or motion masquerade, promoting misclassification (as something other than prey), or a combination of the two.[77]
Comparison of motion camouflage and classical pursuit
Most forms of camouflage are ineffective when the camouflaged animal or object moves, because the motion is easily seen by the observing predator, prey or enemy.[78] However, insects such ashoverflies[79] anddragonflies usemotion camouflage: the hoverflies to approach possible mates, and the dragonflies to approach rivals when defending territories.[80][81] Motion camouflage is achieved by moving so as to stay on a straight line between the target and a fixed point in the landscape; the pursuer thus appears not to move, but only toloom larger in the target's field of vision.[82]Some insects sway while moving to appear to be blown back and forth by the breeze.
The same method can be used for military purposes, for example by missiles to minimise their risk of detection by an enemy.[79] However, missile engineers, and animals such as bats, use the method mainly for its efficiency rather than camouflage.[83]
MaleSyritta pipiens hoverflies use motion camouflage to approach females
MaleAustralian Emperor dragonflies use motion camouflage to approach rivals.
Inmimesis (also calledmasquerade), the camouflaged object looks like something else which is of no special interest to the observer.[84] Mimesis is common inprey animals, for example when apeppered moth caterpillar mimics a twig, or a grasshopper mimics a dry leaf.[85] It is also found in nest structures; some eusocial wasps, such asLeipomeles dorsata, build a nest envelope in patterns that mimic the leaves surrounding the nest.[86]
Mimesis is also employed by somepredators andparasites to lure their prey. For example, aflower mantis mimics a particular kind of flower, such as anorchid.[87] This tactic has occasionally been used in warfare, for example with heavily armedQ-ships disguised as merchant ships.[88][89][90]
Thecommon cuckoo, abrood parasite, provides examples of mimesis both in the adult and in the egg. The female lays her eggs in nests of other, smaller species of bird, one per nest. The female mimics asparrowhawk. The resemblance is sufficient to make small birds take action to avoid the apparent predator. The female cuckoo then has time to lay her egg in their nest without being seen to do so.[91] The cuckoo's egg itself mimics the eggs of the host species, reducing its chance of being rejected.[92][93]
Thezebra's bold pattern may induce motion dazzle in observers
Most forms of camouflage are made ineffective by movement: a deer or grasshopper may be highly cryptic when motionless, but instantly seen when it moves. But one method, motion dazzle, requires rapidly moving bold patterns of contrasting stripes.[94] Motion dazzle may degrade predators' ability to estimate the prey's speed and direction accurately, giving the prey an improved chance of escape.[95] Motion dazzle distorts speed perception and is most effective at high speeds; stripes can also distort perception of size (and so, perceived range to the target). As of 2011, motion dazzle had been proposed for military vehicles, but never applied.[94] Since motion dazzle patterns would make animals more difficult to locate accurately when moving, but easier to see when stationary, there would be an evolutionary trade-off between motion dazzle and crypsis.[95]
An animal that is commonly thought to be dazzle-patterned is thezebra. The bold stripes of the zebra have been claimed to be disruptive camouflage,[96] background-blending and countershading.[97][e] After many years in which the purpose of the coloration was disputed,[98] an experimental study byTim Caro suggested in 2012 that the pattern reduces the attractiveness of stationary models to biting flies such ashorseflies andtsetse flies.[99][100] However, a simulation study by Martin How and Johannes Zanker in 2014 suggests that when moving, the stripes may confuse observers, such as mammalian predators and biting insects, by twovisual illusions: thewagon-wheel effect, where the perceived motion is inverted, and thebarberpole illusion, where the perceived motion is in a wrong direction.[101]
Animals can camouflage themselves by one or more principles using a variety of mechanisms. For example, some animals achieve background matching by changing their skin coloration to resemble their current background.[102]
Four frames of the samepeacock flounder taken a few minutes apart, showing its ability to match its coloration to the environment
Fish and frog melanophore cells change colour by moving pigment-containing bodies.
Each chromatophore contains pigment of only one colour. In fish and frogs, colour change is mediated by a type of chromatophore known asmelanophores that contain dark pigment. A melanophore is star-shaped; it contains many small pigmentedorganelles which can be dispersed throughout the cell, or aggregated near its centre. When the pigmented organelles are dispersed, the cell makes a patch of the animal's skin appear dark; when they are aggregated, most of the cell, and the animal's skin, appears light. In frogs, the change is controlled relatively slowly, mainly byhormones. In fish, the change is controlled by the brain, which sends signals directly to the chromatophores, as well as producing hormones.[105]
The skins of cephalopods such as the octopus contain complex units, each consisting of a chromatophore with surrounding muscle and nerve cells.[106] The cephalopod chromatophore has all its pigment grains in a small elastic sac, which can be stretched or allowed to relax under the control of the brain to vary its opacity. By controlling chromatophores of different colours, cephalopods can rapidly change their skin patterns and colours.[107][108]
On a longer timescale, animals like theArctic hare,Arctic fox,stoat, androck ptarmigan havesnow camouflage, changing their coat colour (by moulting and growing new fur or feathers) from brown or grey in the summer to white in the winter; the Arctic fox is the only species in thedog family to do so.[109] However, Arctic hares which live in the far north ofCanada, where summer is very short, remain white year-round.[109][110]
The principle of varying coloration either rapidly or with the changing seasons has military applications.Active camouflage could in theory make use of both dynamic colour change and counterillumination. Simple methods such as changing uniforms and repainting vehicles for winter have been in use since World War II. In 2011,BAE Systems announced theirAdaptiv infrared camouflage technology. It uses about 1,000 hexagonal panels to cover the sides of a tank. ThePeltier plate panels are heated and cooled to match either the vehicle's surroundings (crypsis), or an object such as a car (mimesis), when viewed in infrared.[111][112][113]
Rock ptarmigan, changing colour in springtime. The male is still mostly in winter plumage
Norwegian volunteer soldiers inWinter War, 1940, with white camouflage overalls over their uniforms
Arctic hares in the low arctic change from brown to white in winter
Snow-camouflaged GermanMarder IIIjagdpanzer and white-overalled crew and infantry in Russia, 1943
Veiled chameleon,Chamaeleo calyptratus, changes colour mainly in relation to mood and for signalling.
Adaptiv infrared camouflage lets an armoured vehicle mimic a car.
Some animals actively seek to hide by decorating themselves with materials such as twigs, sand, or pieces of shell from their environment, to break up their outlines, to conceal the features of their bodies, and to match their backgrounds. For example, acaddisfly larva builds a decorated case and lives almost entirely inside it; adecorator crab covers its back with seaweed, sponges, and stones.[114] Thenymph of the predatorymasked bug uses its hind legs and a 'tarsal fan' to decorate its body with sand or dust. There are two layers of bristles (trichomes) over the body. On these, the nymph spreads an inner layer of fine particles and an outer layer of coarser particles. The camouflage may conceal the bug from both predators and prey.[115][116]
Similar principles can be applied for military purposes, for instance when asniper wears aghillie suit designed to be further camouflaged by decoration with materials such as tufts of grass from the sniper's immediate environment. Such suits were used as early as 1916, the British army having adopted "coats of motley hue and stripes of paint" for snipers.[117] Cott takes the example of the larva of theblotched emerald moth, which fixes a screen of fragments of leaves to its specially hooked bristles, to argue that military camouflage uses the same method, pointing out that the "device is ... essentially the same as one widely practised during the Great War for the concealment, not of caterpillars, but of caterpillar-tractors, [gun] battery positions, observation posts and so forth."[118][119]
Many animals of the open sea, like thisAurelia labiata jellyfish, are largely transparent.
Manymarine animals that float near the surface are highlytransparent, giving them almost perfect camouflage.[120] However, transparency is difficult for bodies made of materials that have differentrefractive indices from seawater. Some marine animals such asjellyfish have gelatinous bodies, composed mainly of water; their thickmesogloea is acellular and highly transparent. This conveniently makes thembuoyant, but it also makes them large for their muscle mass, so they cannot swim fast, making this form of camouflage a costly trade-off with mobility.[120] Gelatinousplanktonic animals are between 50 and 90 percent transparent. A transparency of 50 percent is enough to make an animal invisible to a predator such ascod at a depth of 650 metres (2,130 ft); better transparency is required for invisibility in shallower water, where the light is brighter and predators can see better. For example, a cod can see prey that are 98 percent transparent in optimal lighting in shallow water. Therefore, sufficient transparency for camouflage is more easily achieved in deeper waters.[120]
Some tissues such asmuscles can be made transparent, provided either they are very thin or organised as regular layers or fibrils that are small compared to the wavelength of visible light. A familiar example is the transparency of the lens of the vertebrateeye, which is made of the proteincrystallin, and the vertebratecornea which is made of the proteincollagen.[120] Other structures cannot be made transparent, notably theretinas or equivalent light-absorbing structures of eyes – they must absorb light to be able to function. Thecamera-type eye of vertebrates and cephalopods must be completely opaque.[120] Finally, some structures are visible for a reason, such as to lure prey. For example, thenematocysts (stinging cells) of the transparentsiphonophoreAgalma okenii resemble smallcopepods.[120] Examples of transparent marine animals include a wide variety oflarvae, includingradiata (coelenterates), siphonophores,salps (floatingtunicates),gastropod molluscs,polychaete worms, many shrimplikecrustaceans, and fish; whereas the adults of most of these are opaque and pigmented, resembling the seabed or shores where they live.[120][121] Adultcomb jellies and jellyfish obey the rule, often being mainly transparent. Cott suggests this follows the more general rule that animals resemble their background: in a transparent medium like seawater, that means being transparent.[121] The smallAmazon River fishMicrophilypnus amazonicus and the shrimps it associates with,Pseudopalaemon gouldingi, are so transparent as to be "almost invisible"; further, these species appear to select whether to be transparent or more conventionally mottled (disruptively patterned) according to the local background in the environment.[122]
The adult herring,Clupea harengus, is a typical silvered fish of medium depths, camouflaged by reflection.The herring's reflectors are nearly vertical for camouflage from the side.
Where transparency cannot be achieved, it can be imitated effectively 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. Most fish in the upper ocean such assardine andherring are camouflaged by silvering.[123]
Themarine hatchetfish is extremely flattened laterally, leaving the body just millimetres thick, and the body is so silvery as to resemblealuminium foil. The mirrors consist of microscopic structures similar to those used to providestructural coloration: stacks of between 5 and 10 crystals ofguanine spaced about1⁄4 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.[123]
In fish such as the herring which live 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.[123] Silvering is found in other marine animals as well as fish. Thecephalopods, including squid, octopus and cuttlefish, have multilayer mirrors made of protein rather than guanine.[123]
Counter-illumination means producing light to match a background that is brighter than an animal's body or military vehicle; it is a form of active camouflage. It is notably used by some species ofsquid, such as thefirefly squid and themidwater squid. The latter has light-producing organs (photophores) scattered all over its underside; these create a sparkling glow that prevents the animal from appearing as a dark shape when seen from below.[124] Counterillumination camouflage is the likely function of thebioluminescence of many marine organisms, though light is also produced to attract[125] or to detect prey[126] and for signalling.
Counterillumination has rarely been used for military purposes. "Diffused lighting camouflage" was trialled by Canada'sNational Research Council during the Second World War. It involved projecting light on to the sides of ships to match the faint glow of the night sky, requiring awkward external platforms to support the lamps.[127] The Canadian concept was refined in the AmericanYehudi lights project, and trialled in aircraft includingB-24 Liberators and navalAvengers.[128] The planes were fitted with forward-pointing lamps automatically adjusted to match the brightness of the night sky.[127] This enabled them to approach much closer to a target – within 3,000 yards (2,700 m) – before being seen.[128] Counterillumination was made obsolete byradar, and neither diffused lighting camouflage nor Yehudi lights entered active service.[127]
Blackdevil anglerfish is one of several deep-sea fishes camouflaged against very dark water with a black dermis.
Some deep sea fishes have very black skin, reflecting under 0.5% of ambient light. This can prevent detection by predators or prey fish which use bioluminescence for illumination.Oneirodes had a particularly black skin which reflected only 0.044% of 480 nm wavelength light. The ultra-blackness is achieved with a thin but continuous layer of particles in thedermis,melanosomes. These particles both absorb most of the light, and are sized and shaped so as to scatter rather than reflect most of the rest. Modelling suggests that this camouflage should reduce the distance at which such a fish can be seen by a factor of 6 compared to a fish with a nominal 2% reflectance. Species with this adaptation are widely dispersed in various orders of thephylogenetic tree of bony fishes (Actinopterygii), implying thatnatural selection has driven theconvergent evolution of ultra-blackness camouflage independently many times.[129]
Roman ships, depicted on a 3rd-century ADsarcophagus
Ship camouflage was occasionally used in ancient times.Philostratus (c. 172–250 AD) wrote in hisImagines that Mediterranean pirate ships could be painted blue-gray for concealment.[130]Vegetius (c. 360–400 AD) says that "Venetian blue" (sea green) was used in theGallic Wars, whenJulius Caesar sent hisspeculatoria navigia (reconnaissance boats) to gather intelligence along the coast of Britain; the ships were painted entirely in bluish-green wax, with sails, ropes and crew the same colour.[131] There is little evidence of military use of camouflage on land before 1800, but two unusual ceramics show men inPeru'sMochica culture from before 500 AD, hunting birds with blowpipes which are fitted with a kind of shield near the mouth, perhaps to conceal the hunters' hands and faces.[132] Another early source is a 15th-century French manuscript,The Hunting Book of Gaston Phebus, showing a horse pulling a cart which contains a hunter armed with a crossbow under a cover of branches, perhaps serving as a hide for shooting game.[133]Jamaican Maroons are said to have used plant materials as camouflage in theFirst Maroon War (c. 1655–1740).[134]
The development of military camouflage was driven by the increasing range and accuracy of infantry firearms in the 19th century. In particular the replacement of the inaccuratemusket with weapons such as theBaker rifle made personal concealment in battle essential. TwoNapoleonic War skirmishing units of theBritish Army, the95th Rifle Regiment and the 60th Rifle Regiment, were the first to adopt camouflage in the form of arifle green jacket, while the Line regiments continued to wear scarlet tunics.[135] A contemporary study in 1800 by the English artist and soldierCharles Hamilton Smith provided evidence that grey uniforms were less visible than green ones at a range of 150 yards.[136]
In theAmerican Civil War, rifle units such as the 1st United States Sharp Shooters (in theFederal army) similarly wore green jackets while other units wore more conspicuous colours.[137] The first British Army unit to adoptkhaki uniforms was theCorps of Guides atPeshawar, whenSir Harry Lumsden and his second in command,William Hodson introduced a "drab" uniform in 1848.[138] Hodson wrote that it would be more appropriate for the hot climate, and help make his troops "invisible in a land of dust".[139] Later they improvised by dyeing cloth locally. Other regiments in India soon adopted the khaki uniform, and by 1896khaki drill uniform was used everywhere outside Europe;[140] by theSecond Boer War six years later it was used throughout the British Army.[141]
During the late 19th century camouflage was applied to British coastal fortifications.[142] The fortifications around Plymouth, England were painted in the late 1880s in "irregular patches of red, brown, yellow and green."[143] From 1891 onwards British coastal artillery was permitted to be painted in suitable colours "to harmonise with the surroundings"[144] and by 1904 it was standard practice that artillery and mountings should be painted with "large irregular patches of different colours selected to suit local conditions."[145]
Iron observation post camouflaged as a tree byCubist painterAndré Mare, 1916
In theFirst World War, the French army formed a camouflage corps, led byLucien-Victor Guirand de Scévola,[146][147] employing artists known ascamoufleurs to create schemes such as tree observation posts and covers for guns. Other armies soon followed them.[148][149][150] The termcamouflage probably comes fromcamoufler, aParisian slang term meaningto disguise, and may have been influenced bycamouflet, aFrench term meaningsmoke blown in someone's face.[151][152] The English zoologistJohn Graham Kerr, artist Solomon J. Solomon and the American artist Abbott Thayer led attempts to introduce scientific principles of countershading and disruptive patterning into military camouflage, with limited success.[153][154] In early 1916 theRoyal Naval Air Service began to create dummy air fields to draw the attention of enemy planes to empty land. They created decoy homes and lined fake runways with flares, which were meant to help protect real towns from night raids. This strategy was not common practice and did not succeed at first, but in 1918 it caught the Germans off guard multiple times.[155]
Ship camouflage was introduced in the early 20th century as the range of naval guns increased, with ships painted grey all over.[156][157] In April 1917, when GermanU-boats were sinking many British ships with torpedoes, the marine artistNorman Wilkinson deviseddazzle camouflage, which paradoxically made ships more visible but harder to target.[158] In Wilkinson's own words, dazzle was designed "not for low visibility, but in such a way as to break up her form and thus confuse a submarine officer as to the course on which she was heading".[159]
USSWest Mahomet in dazzle camouflage
Siege howitzer camouflaged against observation from the air, 1917
Austro-Hungarian ski patrol in two-part snow uniforms with improvised head camouflage on Italian front, 1915–1918
In theSecond World War, the zoologist Hugh Cott, aprotégé of Kerr, worked to persuade the British army to use more effective camouflage methods, including countershading, but, like Kerr and Thayer in the First World War, with limited success. For example, he painted two rail-mounted coastal guns, one in conventional style, onecountershaded. In aerial photographs, the countershaded gun was essentially invisible.[160] The power of aerial observation and attack led every warring nation to camouflage targets of all types. TheSoviet Union'sRed Army created the comprehensivedoctrine ofMaskirovka for military deception, including the use of camouflage.[161] For example, during theBattle of Kursk,General Katukov, the commander of the Soviet 1st Tank Army, remarked that the enemy "did not suspect that our well-camouflaged tanks were waiting for him. As we later learned from prisoners, we had managed to move our tanks forward unnoticed". The tanks were concealed in previously prepared defensive emplacements, with only their turrets above ground level.[162] In the air, Second World War fighters were often painted in ground colours above and sky colours below, attempting two different camouflage schemes for observers above and below.[163] Bombers and night fighters were often black,[164] while maritime reconnaissance planes were usually white, to avoid appearing as dark shapes against the sky.[165] For ships, dazzle camouflage was mainly replaced with plain grey in the Second World War, though experimentation with colour schemes continued.[156]
Camouflage has been used to protect military equipment such as vehicles, guns,ships,[156]aircraft and buildings[172] as well as individual soldiers and their positions.[173]Vehicle camouflage methods begin with paint, which offers at best only limited effectiveness. Other methods for stationary land vehicles include covering with improvised materials such as blankets and vegetation, and erecting nets, screens and soft covers which may suitably reflect, scatter or absorbnear infrared andradar waves.[174][175][176] Some military textiles and vehicle camouflage paints also reflect infrared to help provide concealment fromnight vision devices.[177]After the Second World War, radar made camouflage generally less effective, though coastal boats are sometimes painted like land vehicles.[156]Aircraft camouflage too came to be seen as less important because of radar, and aircraft of different air forces, such as the Royal Air Force'sLightning, were often uncamouflaged.[178]
Manycamouflaged textile patterns have been developed to suit the need to matchcombat clothing to different kinds of terrain (such as woodland, snow, and desert).[179] The design of a pattern effective in all terrains has proved elusive.[180][181][182] The AmericanUniversal Camouflage Pattern of 2004 attempted to suit all environments, but was withdrawn after a few years of service.[183] Terrain-specific patterns have sometimes been developed but are ineffective in other terrains.[184] The problem of making a pattern that works at different ranges has been solved with multiscale designs, often with a pixellated appearance and designed digitally, that provide afractal-like range of patch sizes so they appear disruptively coloured both at close range and at a distance.[185] The first genuinely digital camouflage pattern was the Canadian Disruptive Pattern (CADPAT), issued to the army in 2002, soon followed by the American Marine pattern (MARPAT). A pixellated appearance is not essential for this effect, though it is simpler to design and to print.[186]
Hunters of game have long made use of camouflage in the form of materials such as animal skins, mud, foliage, and green or brown clothing to enable them to approach wary game animals.[187]Field sports such asdriven grouse shooting conceal hunters inhides (also called blinds or shooting butts).[188] Modern hunting clothing makes use of fabrics that provide a disruptive camouflage pattern; for example, in 1986 the hunter Bill Jordan created cryptic clothing for hunters, printed with images of specific kinds of vegetation such as grass and branches.[189]
Camouflage is occasionally used to make built structures less conspicuous: for example, inSouth Africa, towers carrying cell telephone antennae are sometimes camouflaged as tall trees with plastic branches, in response to "resistance from the community". Since this method is costly (a figure of three times the normal cost is mentioned), alternative forms of camouflage can include using neutral colours or familiar shapes such as cylinders and flagpoles. Conspicuousness can also be reduced by siting masts near, or on, other structures.[190]
Automotive manufacturers often use patterns to disguise upcoming products. This camouflage is designed to obfuscate the vehicle's visual lines, and is used along with padding, covers, and decals. The patterns' purpose is to prevent visual observation (and to a lesser degree photography), that would subsequently enable reproduction of the vehicle's form factors.[191]
The "dazzle ball" held by the Chelsea Arts Club, 1919
Military camouflage patterns influencedfashion andart from the time of the First World War onwards.Gertrude Stein recalled thecubist artistPablo Picasso's reaction in around 1915:
I very well remember at the beginning of the war being with Picasso on the boulevard Raspail when the first camouflaged truck passed. It was at night, we had heard of camouflage but we had not seen it and Picasso amazed looked at it and then cried out, yes it is we who made it, that is cubism.
In 1919, the attendants of a "dazzle ball", hosted by the Chelsea Arts Club, wore dazzle-patterned black and white clothing. The ball influenced fashion and art via postcards and magazine articles.[193] TheIllustrated London News announced:[193][194]
The scheme of decoration for the great fancy dress ball given by the Chelsea Arts Club at the Albert Hall, the other day, was based on the principles of "Dazzle", the method of "camouflage" used during the war in the painting of ships ... The total effect was brilliant and fantastic.
More recently, fashion designers have often used camouflage fabric for its striking designs, its "patterned disorder" and its symbolism.[195] Camouflage clothing can be worn largely for its symbolic significance rather than for fashion, as when, during the late 1960s and early 1970s in the United States,anti-war protestors often ironically wore military clothing during demonstrations against the American involvement in the Vietnam War.[196]
Modern artists such asIan Hamilton Finlay have used camouflage to reflect on war. His 1973 screenprint of a tank camouflaged in a leaf pattern,Arcadia,[f] is described by theTate as drawing "an ironic parallel between this idea of a natural paradise and the camouflage patterns on a tank".[197] The title refers to theUtopianArcadia of poetry and art, and thememento moriLatin phraseEt in Arcadia ego which recurs in Hamilton Finlay's work. Inscience fiction,Camouflage is a novel aboutshapeshifting alien beings byJoe Haldeman.[198] The word is used more figuratively in works of literature such as Thaisa Frank's collection of stories of love and loss,A Brief History of Camouflage.[199]
In 1986,Andy Warhol began a series of monumental camouflage paintings, which helped to transform camouflage into a popular print pattern. A year later, in 1987, New York designerStephen Sprouse used Warhol's camouflage prints as the basis for his Autumn Winter 1987 collection.[200]
^A letter fromAlfred Russel Wallace to Darwin of 8 March 1868 mentioned such colour change: "Would you like to see the specimens of pupæ of butterflies whose colours have changed in accordance with the colour of the surrounding objects? They are very curious, and Mr.T. W. Wood, who bred them, would, I am sure, be delighted to bring them to show you."[5]
^Cott explains Beddard's observation as a coincident disruptive pattern.[9]
^Before 1860, unpolluted tree trunks were often covered in palelichens; polluted trunks were bare, and often nearly black.
^These distraction markings are sometimes called dazzle markings, but have nothing to do with motion dazzle or wartime dazzle painting.
^The belly of the zebra is white, and the dark stripes narrow towards the belly, so the animal is certainly countershaded, but this does not prove that the main function of the stripes is camouflage.
^Werneck, Jane Margaret Costa de Frontin; Torres, Lucas; Provance, David Willian; Brugnera, Ricardo; Grazia, Jocelia (3 December 2021). "First Report of Predation by a Stink Bug on a Walking-Stick Insect with Reflections on Evolutionary Mechanisms for Camouflage".doi:10.21203/rs.2.10812/v1.S2CID240967012.{{cite journal}}:Cite journal requires|journal= (help)
^Conner, William E. (2014). "Adaptive Sounds and Silences: Acoustic Anti-Predator Strategies in Insects".Insect Hearing and Acoustic Communication. Animal Signals and Communication. Vol. 1. pp. 65–79.doi:10.1007/978-3-642-40462-7_5.ISBN978-3-642-40461-0.ISSN2197-7305.adaptive silence, acoustic crypsis, stealth,
^Miller, Ashadee Kay; Maritz, Bryan; McKay, Shannon; Glaudas, Xavier; Alexander, Graham J. (22 December 2015)."An ambusher's arsenal: chemical crypsis in the puff adder (Bitis arietans)".Proceedings of the Royal Society B: Biological Sciences.282 (1821). The Royal Society: 20152182.doi:10.1098/rspb.2015.2182.ISSN0962-8452.PMC4707760.PMID26674950.Field observations of puff adders (Bitis arietans) going undetected by several scent-orientated predator and prey species led us to investigate chemical crypsis in this ambushing species. We trained dogs (Canis familiaris) and meerkats (Suricata suricatta) to test whether a canid and a herpestid predator could detectB. arietans using olfaction.
^Roosevelt, Theodore (1911). "Revealing and concealing coloration in birds and mammals".Bulletin of the American Museum of Natural History.30 (Article 8):119–231.hdl:2246/470. Roosevelt attacks Thayer on page 191, arguing that neither zebra nor giraffe are "'adequately obliterated' by countershading or coloration pattern or anything else."
^Lev-Yadun, Simcha (2006). "Defensive coloration in plants: a review of current ideas about anti-herbivore coloration strategies". In Teixeira da Silva, J.A. (ed.).Floriculture, ornamental and plant biotechnology: advances and topical issues. Vol. IV. Global Science Books. pp. 292–299.ISBN978-4903313092.
^abKiltie, Richard A. (January 1998). "Countershading: Universally deceptive or deceptively universal?".Trends in Ecology & Evolution.3 (1):21–23.doi:10.1016/0169-5347(88)90079-1.PMID21227055.
^Moskát, C; Honza, M. (2002). "European Cuckoo Cuculus canorus parasitism and host's rejection behaviour in a heavily parasitized Great Reed Warbler Acrocephalus arundinaceus population".Ibis.144 (4):614–622.doi:10.1046/j.1474-919X.2002.00085.x.
^Waage, J. K. (1981). "How the zebra got its stripes: biting flies as selective agents in the evolution of zebra colouration".J. Entom. Soc. South Africa.44:351–358.
^Cloney, R. A.; Florey, E. (1968). "Ultrastructure of Cephalopod Chromatophore Organs".Zeitschrift für Zellforschung und Mikroskopische Anatomie.89 (2):250–280.doi:10.1007/BF00347297.PMID5700268.S2CID26566732.
^Jett, Stephen C. (March 1991). "Further Information on the Geography of the Blowgun and Its Implications for Early Transoceanic Contacts".Annals of the Association of American Geographers.81 (1):89–102.doi:10.1111/j.1467-8306.1991.tb01681.x.JSTOR2563673.
^Payne-Gallwey, Ralph (1903).The Crossbow. Longmans, Green. p. 11.
^Saunders, Nicholas (2005).The People of the Caribbean: An Encyclopedia of Archaeology and Traditional Culture. ABC-CLIO.
^Haythornthwaite, P. (2002).British Rifleman 1797–1815. Osprey Publishing. p. 20.ISBN978-1841761770.
^Stephenson, Hubert Kirk (1948).Applied Physics, pp. 200, 258. Volume 6 of Science in World War II; Office of Scientific Research and Development. Editors: Chauncey Guy Suits and George Russell Harrison. Little, Brown.
^Tinbergen, Niko (1953).The Herring Gull's World. Collins. p. 14.ISBN978-0-00-219444-0.white has proved to be the most efficient concealing coloration for aircraft on anti-submarine patrol{{cite book}}:ISBN / Date incompatibility (help)
^Mellor, D. P. (1958).The Role of Science and Industry. Australia in the War of 1939–1945. Series 4 – Civil. Vol. 5. Canberra: Australian War Memorial. p. 538ff.
^Shapiro, Danielle (2016).John Vassos: Industrial Design for Modern Life. University of Minnesota Press. p. 183.ISBN978-0-8166-9341-2.
^"FM 21–75"(PDF).Chapter 5: Cover, Concealment, and Camouflage. Department of the Army. Archived fromthe original(PDF) on 26 September 2021. Retrieved11 June 2023.
^"FM 21-305/AFMAN 24-306"(PDF).Chapter 20: Vehicle Camouflage And Nuclear, Biological, And Chemical Operations. Department of the Army. pp. 1–9.Archived(PDF) from the original on 17 November 2015. Retrieved16 June 2012.
^"5–103".Appendix D: Camouflage. Department of the Army. Retrieved17 June 2012.
^"SSZ Camouflage". Military Suppliers & News. 2012. Retrieved17 June 2012.
^Jukkola, E. E.; Cohen, R. (1946). "Color Stability of Olive Drab Infrared-Reflecting Camouflage Finishes".Industrial & Engineering Chemistry.38 (9):927–930.doi:10.1021/ie50441a019.
^"The Great Dazzle Ball at the Albert Hall: The Shower of Bomb Balloons and Some Typical Costumes".Illustrated London News. No. 154. 22 March 1919. pp. 414–415.
^"Love and War: The Weaponized Woman".John Galliano for Christian Dior, silk camouflage evening dress. The Museum at FIT. 9 September – 16 December 2006. Archived fromthe original on 12 December 2012. Retrieved1 December 2011.
Newman, Alex; Blechman, Hardy (2004).DPM – Disruptive Pattern Material: An Encyclopaedia of Camouflage: Nature, Military and Culture. DPM.ISBN978-0-9543404-0-7.
Shell, Hanna Rose (2012).Hide and Seek: Camouflage, Photography and the Media of Reconnaissance. Zone Books.ISBN978-1-935-40822-2.
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