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Disruptive coloration (also known as disruptive camouflage or disruptive patterning) is a form of
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
that works by breaking up the outlines of an animal, soldier or
military A military, also known collectively as armed forces, is a heavily armed, highly organized force primarily intended for warfare. It is typically authorized and maintained by a sovereign state, with its members identifiable by their distin ...
vehicle with a strongly contrasting pattern. It is often combined with other methods of
crypsis In ecology, crypsis is the ability of an animal or a plant to avoid observation or detection by other animals. It may be a predation strategy or an antipredator adaptation. Methods include camouflage, nocturnality, subterranean lifestyle an ...
including background colour matching and
countershading Countershading, or Thayer's law, is a method of camouflage in which an animal's coloration is darker on the top or upper side and lighter on the underside of the body. This pattern is found in many species of mammals, reptiles, birds, fish, ...
; special cases are
coincident disruptive coloration Coincident disruptive coloration or coincident disruptive patterns are patterns of disruptive coloration in animals that go beyond the usual camouflage function of breaking up the continuity of an animal's shape, to join up parts of the body that ...
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 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 Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and d ...
Hugh Cott Hugh Bamford Cott (6 July 1900 – 18 April 1987) was a British zoologist, an authority on both natural and military camouflage, and a scientific illustrator and photographer. Many of his field studies took place in Africa, where he was especia ...
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 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 Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
) use patterns that emphasize rather than disrupt their outlines. For example, skunks,
salamanders Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten ...
and
monarch butterflies The monarch butterfly or simply monarch (''Danaus plexippus'') is a milkweed butterfly ( subfamily Danainae) in the family Nymphalidae. Other common names, depending on region, include milkweed, common tiger, wanderer, and black-veined brown. ...
all have high-contrast patterns that display their outlines.


Early research

The artist
Abbott Handerson Thayer Abbott Handerson Thayer (August 12, 1849May 29, 1921) was an American artist, naturalist and teacher. As a painter of portraits, figures, animals and landscapes, he enjoyed a certain prominence during his lifetime, and his paintings are represent ...
in his 1909 book ''
Concealing-Coloration in the Animal Kingdom ''Concealing-Coloration in the Animal Kingdom: An Exposition of the Laws of Disguise Through Color and Pattern; Being a Summary of Abbott H. Thayer’s Discoveries'' is a book published ostensibly by Gerald H. Thayer in 1909, and revised in 191 ...
'' argued that animals were concealed by a combination of
countershading Countershading, or Thayer's law, is a method of camouflage in which an animal's coloration is darker on the top or upper side and lighter on the underside of the body. This pattern is found in many species of mammals, reptiles, birds, fish, ...
and "ruptive" marks, which together "obliterated" their self-shadowing and their shape. Thayer explained that:
Hugh Cott Hugh Bamford Cott (6 July 1900 – 18 April 1987) was a British zoologist, an authority on both natural and military camouflage, and a scientific illustrator and photographer. Many of his field studies took place in Africa, where he was especia ...
'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 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: Further, Cott criticises unscientific attempts at camouflage, early in the
Second World War World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposi ...
, for not understanding the principles involved: 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 Alfred Russel Wallace (8 January 1823 – 7 November 1913) was a British naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution through natural s ...
(''
Darwinism Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations ...
'', 1889),
Edward Bagnall Poulton Sir Edward Bagnall Poulton, FRS HFRSE FLS (27 January 1856 – 20 November 1943) was a British evolutionary biologist, a lifelong advocate of natural selection through a period in which many scientists such as Reginald Punnett doubted it ...
(''
The Colours of Animals ''The Colours of Animals'' is a zoology book written in 1890 by Sir Edward Bagnall Poulton (1856–1943). It was the first substantial textbook to argue the case for Darwinian selection applying to all aspects of animal coloration. The book a ...
'', 1890) and
Frank Evers Beddard Frank Evers Beddard FRS FRSE (19 June 1858 – 14 July 1925) was an English zoologist. He became a leading authority on annelids, including earthworms. He won the Linnean Medal in 1916 for his book on oligochaetes. Life Beddard was born in ...
(''
Animal Coloration Animal coloration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male ...
'', 1895); Stevens observes that background matching on its own would always fail because of "discontinuities between the boundary of the animal and the background".


In animals

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 The leopard (''Panthera pardus'') is one of the five extant species in the genus ''Panthera'', a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia ...
, and some potential prey like the
Egyptian nightjar The Egyptian nightjar (''Caprimulgus aegyptius'') is a medium-small nightjar which occurs in south west Asia and north Africa and winters in tropical Africa. This is a fairly common species with a wide distribution which faces no obvious threats ...
, use disruptive patterns. Disruptive patterns are defined by A. Barbosa and colleagues as "characterized by high-contrast light and dark patches, in 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 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 the lightness of the background
oak An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably ...
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 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, 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.


The opposite case: aposematism

Many poisonous or distasteful animals that advertise their presence with warning coloration (
aposematism Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
) use patterns that emphasize rather than disrupt their outlines. For example, skunks,
salamanders Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten ...
and
monarch butterflies The monarch butterfly or simply monarch (''Danaus plexippus'') is a milkweed butterfly ( subfamily Danainae) in the family Nymphalidae. Other common names, depending on region, include milkweed, common tiger, wanderer, and black-veined brown. ...
all have high contrast patterns that display their outlines. These advertising patterns exploit 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.


A disputed case: the giraffe

The presence of bold markings does not in itself prove that an animal relies on camouflage. Roosevelt attacks Thayer on page 191, arguing that neither zebra nor giraffe are "'adequately obliterated' by countershading or coloration pattern or anything else." According to Mitchell, adult
giraffe The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, ''Giraffa camelopardali ...
s are "inescapably conspicuous", making 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 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.


Other ways of hiding outlines

The outlines of an animal's body can be made hard to see by other methods, such as by using 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.


In plants

The possibility of protective coloration in plants has been little studied. T. J. Givnish and Simcha Lev-Yadun have proposed that leaf
variegation Variegation is the appearance of differently coloured zones in the leaves and sometimes the stems and fruit of plants. Species with variegated individuals are sometimes found in the understory of tropical rainforests, and this habitat is the s ...
with white spots may serve as camouflage in forest understory plants, where there 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
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthp ...
s; and the hypothesis is testable.


Military usage

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 the 1940s have been disruptively coloured, and with the issue of US Woodland pattern to
United States armed forces The United States Armed Forces are the Military, military forces of the United States. The armed forces consists of six Military branch, service branches: the United States Army, Army, United States Marine Corps, Marine Corps, United States N ...
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 The British Armed Forces, also known as His Majesty's Armed Forces, are the military forces responsible for the defence of the United Kingdom, its Overseas Territories and the Crown Dependencies. They also promote the UK's wider interests, ...
and was later used by many other armies.


Challenges

Three major challenges face the design of disruptively patterned uniforms. Firstly, units frequently move from one terrain to another, where 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 was applied to British vehicles operating in
Berlin Berlin ( , ) is the capital and largest city of Germany by both area and population. Its 3.7 million inhabitants make it the European Union's most populous city, according to population within city limits. One of Germany's sixteen constit ...
during the
Cold War The Cold War is a term commonly used to refer to a period of geopolitical tension between the United States and the Soviet Union and their respective allies, the Western Bloc and the Eastern Bloc. The term '' cold war'' is used because t ...
, 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 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 In mathematics, a fractal is a geometric shape containing detailed structure at arbitrarily small scales, usually having a fractal dimension strictly exceeding the topological dimension. Many fractals appear similar at various scales, as ill ...
-like range of patch sizes, enabling them to be effectively disruptive both at close range and at a distance. The first genuinely digital camouflage pattern was the Canadian CADPAT, soon followed by the American
MARPAT MARPAT (short for Marine pattern) is a multi-scale camouflage pattern in use with the United States Marine Corps, designed in 2001 and introduced from late 2002 to early 2005 with the Marine Corps Combat Utility Uniform (MCCUU), which replace ...
. A pixellated appearance is not essential for this effect, though it is simpler to design and to print.


Examples

File:WoodFrog DarienLakesStatePark 2020-06-16 (02).jpg,
Wood frog ''Lithobates sylvaticus'' or ''Rana sylvatica'', commonly known as the wood frog, is a frog species that has a broad distribution over North America, extending from the boreal forest of the north to the southern Appalachians, with several nota ...
among fallen leaves. File:Great male Leopard in South Afrika-JD.JPG, alt=photo of a leopard,
Leopard The leopard (''Panthera pardus'') is one of the five extant species in the genus ''Panthera'', a member of the cat family, Felidae. It occurs in a wide range in sub-Saharan Africa, in some parts of Western and Central Asia, Southern Russia ...
: a disruptively camouflaged (and
countershaded Countershading, or Thayer's law, is a method of camouflage in which an animal's coloration is darker on the top or upper side and lighter on the underside of the body. This pattern is found in many species of mammals, reptiles, birds, fish, ...
) predator File:Ptarmigan and five chicks.JPG, alt=photo of a hen ptarmigan and her chicks, A
ptarmigan ''Lagopus'' is a small genus of birds in the grouse subfamily commonly known as ptarmigans (). The genus contains three living species with numerous described subspecies, all living in tundra or cold upland areas. Taxonomy and etymology The g ...
and five chicks with exceptional disruptive camouflage File:Jumping spider with prey.jpg, alt=photo of a jumping spider,
Jumping spider Jumping spiders are a group of spiders that constitute the family Salticidae. As of 2019, this family contained over 600 described genera and over 6,000 described species, making it the largest family of spiders at 13% of all species. Jumping spi ...
: a disruptively camouflaged invertebrate predator File:Armoured personnel carriers, Eriboll - geograph.org.uk - 1316295.jpg, alt=photo of armoured vehicles easy to see on bare hillside, "Shape, shine, shadow" make these "camouflaged" military vehicles easily visible, their outlines not disrupted File:Challenger 2 Tank MOD 45148907.jpg, alt=photo of a Challenger 2 tank at speed on grassy plain, A British Challenger 2 tank painted in bold disruptive pattern of sand and green File:Chaetodon striatus Brasil.jpg, The banded butterflyfish, '' Chaetodon striatus'', has strong disruptive bands through the body and concealing the eye. File:US Navy WWII ship camouflage measures - detail of USS Alabama in measure 16.jpg, USS ''Alabama'' wearing Measure 12 ship camouflage during World War II File:A-7D Corsairs 354th TFW at Korat 1972.JPG, Disruptively camouflaged A-7D Corsairs on a disruptively painted concrete surface, Thailand, 1972


References


Sources

* * {{camouflage Camouflage Military camouflage