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71-609: German World War II camouflage patterns formed a family of disruptively patterned military camouflage designs for clothing, used and in the main designed during the Second World War . The first pattern, Splittertarnmuster ("splinter camouflage pattern"), was designed in 1931 and was initially intended for Zeltbahn shelter halves. The clothing patterns developed from it combined a pattern of interlocking irregular green, brown, and buff polygons with vertical "rain" streaks. Later patterns, all said to have been designed for

142-555: A Munich art professor and then the director of the German camouflage research unit, at the request of an SS Major, Wim Brandt. Brandt was an engineer and the commander of the SS-VT reconnaissance battalion, and he was looking for better camouflage. Schick had researched the effect of light on trees in summer and in autumn. These led to the idea of reversible camouflage clothing, with green summer patterns on one side, brown autumn patterns on

213-479: A disruptive pattern is to prevent, or to delay as long as possible, the first recognition of an object by sight... irregular patches of contrasted colours and tones ... tend to catch the eye of the observer and to draw his attention away from the shape which bears them. Further, Cott criticises unscientific attempts at camouflage, early in the Second World War , for not understanding the principles involved: Various recent attempts to camouflage tanks, armoured cars and

284-567: A distance. The first genuinely digital camouflage pattern was the Canadian CADPAT , soon followed by the American MARPAT . A pixellated appearance is not essential for this effect, though it is simpler to design and to print. Countershading 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

355-827: A distinctive emblem of the Waffen-SS during the war. However, patterned uniforms were worn by some other units, including from 1941 the Luftwaffe , which had its own version of Splittertarnmuster , as well as the Kriegsmarine (navy), the Fallschirmjäger (paratroops), and the Waffen-SS. The 1945 Leibermuster was planned to be issued to both the SS and the Wehrmacht, but it appeared too late to be widely distributed. Production of groundsheets, helmet covers and smocks by

426-448: A highly irregular outline. For example, the comma butterfly , Polygonia c-album , is highly cryptic when its wings are closed, with cryptic colours, disruptive pattern, and irregular outer margins to the wings. The possibility of protective coloration in plants has been little studied. T. J. Givnish and Simcha Lev-Yadun have proposed that leaf variegation with white spots may serve as camouflage in forest understory plants, where there

497-492: 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

568-454: A mother does not affect survival, Mitchell suggests that young giraffes must be extremely well camouflaged. This is supported by the fact that coat markings are strongly inherited. Conversely, far from hiding, 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. The outlines of an animal's body can be made hard to see by other methods, such as by using

639-422: A nonrepetitive configuration, that also provide camouflage by disrupting the recognizable shape or orientation of the animal", as in the cuttlefish. The strategy appears paradoxical and counter-intuitive as a method of camouflage, since disrupting outlines depends on using patches of colour which contrast strongly with each other, so the patches are themselves conspicuous. While background matching works best for

710-713: A patent in 1902 to paint warships, both submarines and surface ships , using countershading, but failed to convince the US Navy to adopt his ideas. Hugh Bamford Cott in his 1940 book Adaptive Coloration in Animals described many instances of countershading, following Thayer in general approach 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

781-475: A plain background", but at once adds that conditions are hardly ever ideal, as they are constantly changing, as is the light. Therefore, Cott argues, camouflage has to break up the perceived continuous surfaces of an object and its outlines. In his own words, "for effective concealment, it is essential that the tell-tale appearance of form should be destroyed." He draws an analogy with a pickpocket who carefully distracts your attention, arguing that: The function of

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852-492: 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." Cott's gun

923-655: 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. 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. Despite demonstrations and examples adduced by Cott and others, little experimental evidence for

994-544: A single background, disruptive coloration is a more effective strategy when an animal or a military vehicle may have a variety of backgrounds. Martin Stevens and colleagues in 2006 made what they believed was the first experimental test that "disruptive coloration is effective even when some colour patches do not match the background and have a high contrast with both the background and adjacent pattern elements (disruptive contrast)". They used "moth-like targets", some matching

1065-531: A single background, disruptive coloration is a more effective strategy when an animal or a military vehicle may have a variety of backgrounds. Conversely, poisonous or distasteful animals that advertise their presence with warning coloration ( aposematism ) use patterns that emphasize rather than disrupt their outlines. For example, skunks , salamanders and monarch butterflies all have high-contrast patterns that display their outlines. The artist Abbott Handerson Thayer in his 1909 book Concealing-Coloration in

1136-399: 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

1207-422: Is a dappled background. Lev-Yadun has also suggested, however, that similar markings serve as conspicuous warning coloration in well-defended thorny plants of open habitats, where the background is uniformly bright. Givnish found a correlation of leaf mottling with closed habitats. Disruptive camouflage would have a clear evolutionary advantage in plants: they would tend to escape from being eaten by herbivores ; and

1278-519: Is apparently the basal and predominant use of almost all the bolder patterns in animals' costumes. Hugh Cott 's 1940 book Adaptive Coloration in Animals introduced ideas such as "maximum disruptive contrast". This uses streaks of boldly contrasting colour, which paradoxically make animals or military vehicles less visible by breaking up their outlines. He explains that in ideal conditions, background colour matching together with countershading would "suffice to render an animal absolutely invisible against

1349-531: 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. 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. Rowland (2009) identified an additional mechanism of countershading not previously analysed, namely that

1420-474: The Waffen-SS by Johann Georg Otto Schick , evolved into more leaf-like forms with rounded dots or irregular shapes. Camouflage smocks were designed to be reversible, providing camouflage for two seasons, whether summer and autumn, or summer and winter (snow). Distribution was limited to the Waffen-SS, ostensibly because of a patent, though variants were used by other units, including the Luftwaffe . Production

1491-498: The embryo that will become the belly skin causes the Agouti gene to become active there, creating the countershading seen in adult mammals. 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

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1562-422: The gray snapper , Lutianus griseus , is effectively flattened by its countershading, while it hunts an "almost invisible" prey, the hardhead silverside , Atherina laticeps which swims over greyish sands. 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 . It tones

1633-426: The white which the under surfaces and parts in shade should be painted. 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. Research by Ariel Tankus and Yehezkel Yeshurun investigating "camouflage breaking",

1704-575: The 1940s have been disruptively coloured, and with the issue of US Woodland pattern to United States armed forces from 1981, disruptive pattern became a dominant feature of military uniforms. From 1969, Disruptive Pattern Material (DPM) began to replace plain material for uniforms in the British Armed Forces and was later used by many other armies. Three major challenges face the design of disruptively patterned uniforms. Firstly, units frequently move from one terrain to another, where

1775-683: The Animal Kingdom argued that animals were concealed by a combination of countershading and "ruptive" marks, which together "obliterated" their self-shadowing and their shape. Thayer explained that: Markings... of whatever sort, tend to obliterate ,—to cancel, by their separate and conflicting pattern, the visibility of the details and boundaries of form.... If the bird's or butterfly's costume consists of sharply contrasted bold patterns of light and dark, in about equal proportions, its contour will be "broken up" against both light and dark—light failing to show against light, dark against dark. Such

1846-470: The Middle East , and Kerr unsuccessfully intervened, pleading for guns to be painted Cott's way and Cott to be brought home. 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

1917-608: The Warei, Forster and Joring companies began in November 1938. They were initially hand-printed, limiting deliveries by January 1939 to only 8,400 groundsheets and 6,800 helmet covers and a small number of smocks. By June 1940, machine printing had taken over, and 33,000 smocks were made for the Waffen-SS. Supplies of high quality cotton duck, however, remained critically short throughout the war, and essentially ran out in January 1943. It

1988-524: 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

2059-528: The authors observed that animals with Thayer countershading are using "counter-measures to convexity based detectors", which implied "predators who use convexity based detectors." 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 . She noted there had been "much debate" about how countershading works. She considered

2130-450: 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

2201-408: The background colours and contrasts may differ greatly. A uniform designed for woodland will be too strongly contrasting for desert use, and too green for urban use. Therefore, no single camouflage pattern is effective in all terrains. The American UCP of 2004 attempted to suit all environments but was withdrawn after a few years of service. Terrain specific patterns like "Berlin camouflage", which

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2272-566: The background is bright enough to make an animal that is not counter-illuminated appear as a shadow. 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

2343-462: The belly is partly concealed, but that when the squirrel is vertical (as when climbing a tree trunk) this effect did not occur. Thayer's original argument, restated by Cott, 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

2414-674: 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. The English zoologist Edward Bagnall Poulton , author of The Colours of Animals (1890) discovered

2485-477: 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

2556-490: 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." Countershading is observed in a wide range of animal groups, both terrestrial, such as deer , and marine, such as sharks . It is the basis of camouflage in both predators and prey. It is used alongside other forms of camouflage including colour matching and disruptive coloration. Among predatory fish,

2627-482: The boundary of the animal and the background". Disruptive patterns use strongly contrasting markings such as spots or stripes to break up the outlines of an animal or military vehicle. Some predators, like the leopard , and some potential prey like the Egyptian nightjar , use disruptive patterns. Disruptive patterns are defined by A. Barbosa and colleagues as "characterized by high-contrast light and dark patches, in

2698-583: 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

2769-409: The conclusion that their patterns are for camouflage appear counterintuitive: but when standing among trees and bushes, their camouflage is effective at even a few metres' distance. Further, young giraffes are much more vulnerable to predation than adults: between 60% and 75% of calves die within a year. Mothers hide their calves, which spend much of the time lying down in cover. Since the presence of

2840-407: The countershading of various insects, including the pupa or chrysalis of the purple emperor butterfly, Apatura iris , the caterpillar larvae of the brimstone moth, Opisthograptis luteolata and of the peppered moth, Biston betularia . However he did not use the term countershading, nor did he suggest that the effect occurred widely. The New Hampshire artist Abbott Handerson Thayer

2911-723: The dinosaur was optimally countershaded for a closed habitat such as a forest. Another form of animal camouflage uses bioluminescence to increase the average brightness of an animal to match the brightness of the background. 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. 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

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2982-601: The effect and render the work practically useless. The pioneering work of Thayer and Cott is endorsed in the 2006 review of disruptive coloration by Martin Stevens and colleagues, which notes that they proposed a "different form of camouflage" from the traditional "strategy of background matching" proposed by authors such as Alfred Russel Wallace ( Darwinism , 1889), Edward Bagnall Poulton ( The Colours of Animals , 1890) and Frank Evers Beddard ( Animal Coloration , 1895); Stevens observes that background matching on its own would always fail because of "discontinuities between

3053-562: 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 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. Evolutionary developmental biology has assembled evidence from embryology and genetics to show how evolution has acted at all scales from

3124-464: The effectiveness of countershading, without succeeding in persuading their armed forces to adopt the technique, though they influenced military adoption of camouflage in general. 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

3195-468: The enemy observing the pattern. A pattern printed with small patches of colour blends into a single perceived mass at a certain range, defeating the disruptive effect. Conversely, a pattern printed with large patches of colour appears conspicuous at shorter ranges. This problem has been solved with pixellated shapes, often designed digitally, that provide a fractal -like range of patch sizes, enabling them to be effectively disruptive both at close range and at

3266-622: The evidence for Thayer's theory that this acts as camouflage "by reducing ventral shadowing", and reviewed alternative explanations for countershading. 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". These are examined in turn below. Cott, like Thayer, argued that countershading would make animals hard to see from

3337-452: The eye with a disruptive eye mask , sometimes contrasting with a stripe above the eye , making it seem just part of a dark area of background. Cott called this a special case of a " coincident disruptive pattern ". Another camouflage mechanism, distractive markings , also involves conspicuous marks and has for a century since Thayer's initial description been conflated with it, but the two require different kinds of marking. For distraction,

3408-513: The hypothesis is testable. Disruptive coloration is common in military usage, for military vehicles, for firing positions and other installations, and for individual soldiers, where uniforms, equipment such as helmets, and face paint may be used to break up outlines and features. Disruptive coloration, however, does not always achieve crypsis on its own, as an animal or a military target may be given away by other factors including shape, shine, and shadow. Many military camouflage patterns since

3479-761: The lightness of the background oak tree bark, other mismatching it, each with a dead mealworm. If the mealworm was removed it was assumed a bird predator had taken it: this could be distinguished from visits by other predators. They found that disruptive coloration provided the best protection from bird predators when the pattern was matched to background luminance, but even when elements in a pattern did not match, disruptive patterns were still better at reducing predation than either non-disruptive patterns or plain (unpatterned) control targets. Disruptive patterns can also conceal specific features. Animals such as fish, birds, frogs and snakes can readily be detected by their eyes, which are necessarily round and dark. Many species conceal

3550-513: The markings should be small and should avoid the prey's outline so as to take attention away from it, whereas disruptive markings should contact the outline so as to break it up. Many poisonous or distasteful animals that advertise their presence with warning coloration ( aposematism ) use patterns that emphasize rather than disrupt their outlines. For example, skunks , salamanders and monarch butterflies all have high contrast patterns that display their outlines. These advertising patterns exploit

3621-574: The opposite principle to disruptive coloration, for what is in effect the exactly opposite effect: to make the animal as conspicuous as possible. Some Lepidoptera, including the wood tiger moth , are aposematic and disruptively coloured; against a green, vegetative background their bright aposematic coloration stands out, but on the ground their wings camouflage them among dead leaves and dirt. The presence of bold markings does not in itself prove that an animal relies on camouflage. According to Mitchell, adult giraffes are "inescapably conspicuous", making

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3692-602: The other. In 1937, the patterns were field tested by the SS-VT Deutschland regiment, resulting in an estimate that they would cut casualties by fifteen percent. In 1938, a reversible spring/autumn helmet cover, smock, and sniper's face mask in Schick's forest patterns on waterproof cotton duck were patented for the Waffen-SS. The patent is said to have prevented the Wehrmacht from using the patterns, which became

3763-419: 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

3834-445: The outlines of an animal, soldier or military hardware with a strongly contrasting pattern. It is often combined with other methods of crypsis including background colour matching and countershading ; special cases are coincident disruptive coloration and the disruptive eye mask seen in some fishes, amphibians, and reptiles. It appears paradoxical as a way of not being seen, since disruption of outlines depends on high contrast, so

3905-433: The patches of colour are themselves conspicuous. The importance of high-contrast patterns for successful disruption was predicted in general terms by the artist Abbott Thayer in 1909 and explicitly by the zoologist Hugh Cott in 1940. Later experimental research has started to confirm these predictions. Disruptive patterns work best when all their components match the background. While background matching works best for

3976-490: The roofs of buildings with paint reveal an almost complete failure by those responsible to grasp the essential factor in the disguise of surface continuity and of contour. Such work must be carried out with courage and confidence, for at close range objects properly treated will appear glaringly conspicuous. But they are not painted for deception at close range, but at ranges at which ... bombing raids are likely... And at these distances differences of tint ... blend and thus nullify

4047-417: 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. 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

4118-488: The shoulders and arms of battledress should be countershaded. 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: 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

4189-402: The side, as they would "fade into a ghostly elusiveness". 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." Kiltie measured the effect of the countershading of the eastern gray squirrel , Sciurus carolinensis , showing that when the squirrel is horizontal the self-shadowing of

4260-569: The skunk, and the sharp claws, aggressive nature and stink of the honey badger. 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. 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

4331-619: 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 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. Thayer obtained

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4402-518: The surface waters of the sea, that countershading reaches its maximum development and significance. 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. 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

4473-529: The theory to a fantastic extreme". 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" or in Cott's words "Colour resemblance alone is not sufficient to afford concealment". Cott explained that Contrary to what might have been expected by any one lacking in artistic perception,

4544-538: 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

4615-422: 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. There

4686-451: 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

4757-506: Was applied to British vehicles operating in Berlin during the Cold War , have sometimes been developed but are ineffective in other terrains. Secondly, the effectiveness of any pattern in disrupting a soldier's outlines varies with lighting, depending on the weather and the height of the sun in the sky. And thirdly, any given patch of printed colour varies in apparent size with distance from

4828-594: Was limited by shortage of materials, especially of high quality waterproof cotton duck . The Reichswehr (Army of the Weimar Republic ) started experimenting with camouflage patterns for Wehrmacht uniforms before World War II and some army units used Splittertarnmuster ("splinter camouflage pattern"), first issued in 1931, and based on Zeltbahn shelter halves/groundsheets. Waffen-SS combat units used various patterns from 1935 onwards. The SS camouflage patterns were designed by Johann Georg Otto Schick,

4899-453: 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. Early researchers including Alfred Russel Wallace , Beddard, Cott and Craik argued that in marine animals including pelagic fish such as marlin and mackerel , as well as dolphins , sharks , and penguins

4970-444: 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. 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

5041-410: Was replaced by non-waterproof cotton drill cloth. The German names used for the plane tree, palm and oak leaf patterns are not those that were used in the German armed forces, but were invented by postwar collectors of militaria. Disruptively patterned Disruptive coloration (also known as disruptive camouflage or disruptive patterning ) is a form of camouflage that works by breaking up

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