Camouflage

The peacock flounder can change its pattern and

colours to match its environment.
A soldier applying camouflage face paint; both helmet
and jacket are disruptively patterned.

Camouflage is the use of any combination

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

Military camouflage was spurred by the

increasing range and accuracy of firearms
in the 19th century. In particular the
replacement of the inaccurate musket with
the rifle made personal concealment in
battle a survival skill. In the 20th century,
military camouflage developed rapidly,
especially during the First World War. On
land, artists such as André Mare designed
camouflage schemes and observation
posts disguised as trees. At sea, merchant
ships and troop carriers were painted in
dazzle patterns that were highly visible,
but designed to confuse enemy
submarines as to the target's speed, range,
and heading. During and after the Second
World War, a variety of camouflage
schemes were used for aircraft 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

making cell 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.

History
In zoology

Octopuses like this Octopus cyanea can change colour

(and shape) for camouflage
In ancient Greece, Aristotle (384 BC – 322
BC) commented on the colour-changing
abilities, both for camouflage and for
signalling, of cephalopods including the
octopus, in his Historia animalium:[1]

The octopus ... seeks its prey by so

changing its colour as to render it
like the colour of the stones
adjacent to it; it does so also
when alarmed.

— Aristotle.[1]
Camouflage has been a topic of interest
and research in zoology for well over a
century. According to Charles Darwin's
1859 theory of natural selection,[2] features
such as camouflage evolved by providing
individual animals with a reproductive
advantage, enabling them to leave more
offspring, on average, than other members
of the same species. In his Origin of
Species, Darwin wrote:[3]

When we see leaf-eating insects

green, and bark-feeders mottled-
grey; the alpine ptarmigan white
in winter, the red-grouse the
colour of heather, and the black-
grouse that of peaty 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 from birds 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]

Experiment by Poulton, 1890: swallowtailed moth

pupae camouflaged to match their backgrounds when
larvae
The English zoologist Edward Bagnall
Poulton studied animal coloration,
especially camouflage. In his 1890 book
The 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
that swallowtailed moth pupae were
camouflaged to match the backgrounds
on which they were reared as larvae.[4][a]
Poulton's "general protective
resemblance"[6] was at that time
considered to be the main method of
camouflage, as when Frank Evers Beddard
wrote in 1892 that "tree-frequenting
animals are often green in colour. Among
vertebrates numerous species of parrots,
iguanas, tree-frogs, and the green tree-
snake are examples".[7] Beddard did
however briefly mention other methods,
including the "alluring coloration" of the
flower mantis and the possibility of a
different mechanism in the orange 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 [an umbellifer], so close is their
mutual resemblance."[8][b] He also
explained the coloration of sea fish such
as the mackerel: "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."[10]

Abbott Thayer's 1907 painting Peacock in the Woods

depicted a peacock as if it were camouflaged.
The artist Abbott Handerson Thayer
formulated what is sometimes called
Thayer's Law, the principle of
countershading.[11] However, he
overstated the case in the 1909 book
Concealing-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 as Peacock in the
Woods (1907) to reinforce his
argument.[14] Thayer was roundly mocked
for these views by critics including Teddy
Roosevelt.[15]

The English zoologist Hugh Cott's 1940

book Adaptive 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 how disruptive 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 a natural history narrative which
illustrated theories with examples.[19]
Camouflage is a soft-tissue feature that is
rarely preserved in the fossil record, but
rare fossilised skin samples from the
Cretaceous period show that some marine
reptiles were countershaded. The skins,
pigmented with dark-coloured eumelanin,
reveal that both leatherback turtles and
mosasaurs had dark backs and light
bellies.[20]

Military

Before 1800
Roman ships, depicted on a 3rd-century AD
sarcophagus

Ship camouflage was occasionally used in

ancient times. Philostratus (c. 172–250
AD) wrote in his Imagines that
Mediterranean pirate ships could be
painted blue-gray for
concealment.[21]Vegetius (c. 360–400 AD)
says that "Venetian blue" (sea green) was
used in the Gallic Wars, when Julius
Caesar sent his speculatoria 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.[22] There is little evidence of
military use of camouflage on land before
1800, but two unusual ceramics show men
in Peru's Mochica 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.[23] 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.[24] Jamaican Maroons are
said to have used plant materials as
camouflage in the First Maroon War (c.
1655–1740).[25]

19th-century origins

Green-jacketed rifleman firing Baker rifle 1803

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 inaccurate musket with weapons such
as the Baker rifle made personal
concealment in battle essential. Two
Napoleonic War skirmishing units of the
British Army, the 95th Rifle Regiment and
the 60th Rifle Regiment, were the first to
adopt camouflage in the form of a rifle
green jacket, while the Line regiments
continued to wear scarlet tunics.[26] A
contemporary study in 1800 by the English
artist and soldier Charles Hamilton Smith
provided evidence that grey uniforms were
less visible than green ones at a range of
150 yards.[27]
In the American Civil War, rifle units such
as the 1st United States Sharp Shooters
(in the Federal army) similarly wore green
jackets while other units wore more
conspicuous colours.[28] The first British
Army unit to adopt khaki uniforms was the
Corps of Guides at Peshawar, when Sir
Harry Lumsden and his second in
command, William Hodson introduced a
"drab" uniform in 1848.[29] Hodson wrote
that it would be more appropriate for the
hot climate, and help make his troops
"invisible in a land of dust".[30] Later they
improvised by dyeing cloth locally. Other
regiments in India soon adopted the khaki
uniform, and by 1896 khaki drill uniform
was used everywhere outside Europe;[31]
by the Second Boer War six years later it
was used throughout the British Army.[32]

First World War

Iron observation post camouflaged as a tree by Cubist

painter André Mare, 1916

In the First World War, the French army

formed a camouflage corps, led by Lucien-
Victor Guirand de Scévola,[33][34] employing
artists known as camoufleurs to create
schemes such as tree observation posts
and covers for guns. Other armies soon
followed them.[35][36][37] The term
camouflage probably comes from
camoufler, a Parisian slang term meaning
to disguise, and may have been influenced
by camouflet, a French term meaning
smoke blown in someone's face.[38][39] The
English zoologist John 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.[40][41]

Ship camouflage was introduced in the

early 20th century as the range of naval
guns increased, with ships painted grey all
over.[42][43] In April 1917, when German U-
boats were sinking many British ships with
torpedoes, the marine artist Norman
Wilkinson devised dazzle camouflage,
which paradoxically made ships more
visible but harder to target.[44] 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".[45]

USS West 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

Second World War

In the Second World War, the zoologist

Hugh Cott, a protégé of Kerr, worked to
persuade the British army to use more
effective camouflage techniques, 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, one countershaded. In aerial
photographs, the countershaded gun was
essentially invisible.[46] The power of aerial
observation and attack led every warring
nation to camouflage targets of all types.
The Soviet Union's Red Army created the
comprehensive doctrine of Maskirovka for
military deception, including the use of
camouflage.[47] For example, during the
Battle 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.[48] 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.[49] Bombers and night fighters
were often black,[50] while maritime
reconnaissance planes were usually white,
to avoid appearing as dark shapes against
the sky.[51] For ships, dazzle camouflage
was mainly replaced with plain grey in the
Second World War, though
experimentation with colour schemes
continued.[42]

As in the First World War, artists were

pressed into service; for example, the
surrealist painter Roland Penrose became
a lecturer at the newly founded
Camouflage Development and Training
Centre at Farnham Castle,[52] writing the
practical Home Guard Manual of
Camouflage.[53] The film-maker Geoffrey
Barkas ran the Middle East Command
Camouflage Directorate during the 1941–
1942 war in the Western Desert, including
the successful deception of Operation
Bertram. Hugh Cott was chief instructor;
the artist camouflage officers, who called
themselves camoufleurs, included Steven
Sykes and Tony Ayrton.[54][55] In Australia,
artists were also prominent in the Sydney
Camouflage Group, formed under the
chairmanship of Professor William John
Dakin, a zoologist from Sydney University.
Max Dupain, Sydney Ure Smith and William
Dobell were among the members of the
group, which worked at Bankstown Airport,
RAAF Base Richmond and Garden Island
Dockyard.[56]
Maritime patrol Catalina, painted white to
minimise visibility against the sky

1937 summer variant of Waffen SS

Flecktarn Plane tree pattern
USS Duluth in naval camouflage Measure
32, Design 11a, one of many dazzle
schemes used on warships

A Spitfire's underside 'azure' paint scheme,

meant to hide it against the sky
 A Luftwaffe aircraft hangar built to
resemble a street of village houses,
Belgium, 1944

Red Army soldiers in the Battle of

Stalingrad in snow camouflage overalls,
January 1943

After 1945
Camouflage has been used to protect
military equipment such as vehicles, guns,
ships,[42] aircraft and buildings[57] as well
as individual soldiers and their
positions.[58] Vehicle camouflage
techniques 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
absorb near infrared and radar
waves.[59][60][61] Some military textiles and
vehicle camouflage paints also reflect
infrared to help provide concealment from
night vision devices.[62] After the Second
World War, radar made camouflage
generally less effective, though coastal
boats are sometimes painted like land
vehicles.[42] 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's Lightning,
were often uncamouflaged.[63]

Many camouflaged textile patterns have

been developed to suit the need to match
combat clothing to different kinds of
terrain (such as woodland, snow, and
desert).[64] The design of a pattern
effective in all terrains has proved
elusive.[65][66][67] The American Universal
Camouflage Pattern of 2004 attempted to
suit all environments, but was withdrawn
after a few years of service.[68] Terrain-
specific patterns have sometimes been
developed but are ineffective in other
terrains.[69] The problem of making a
pattern that works at different ranges has
been solved with pixellated shapes, often
designed digitally, that provide a fractal-
like range of patch sizes so they appear
disruptively coloured both at close range
and at a distance.[70] The first genuinely
digital camouflage pattern was the
Canadian CADPAT, issued to the army in
2002, soon followed by the American
MARPAT. A pixellated appearance is not
essential for this effect, though it is
simpler to design and to print.[71][72]
 CADPAT was the first pixellated digital
camouflage pattern to be issued, in 2002.

British Disruptive Pattern Material, issued

to special forces in 1963 and universally
by 1968
 2007 2-colour snow variant of Finnish
Defence Forces M05 pattern

Main (4-colour woodland) variant of

Chinese People's Liberation Army Type 99
pattern, c. 2006
  

Modern German Flecktarn 1990,

developed from a 1938 pattern, a non-
digital pattern which works at different
distances
  

US "Chocolate Chip" Six-Color Desert

Pattern developed in 1962, widely used in
Gulf War

Principles

Draco indochinensis uses several methods of

fl i l di di ti l ti l i fl t
camouflage, including disruptive coloration, lying flat,
and concealment of shadow.

Camouflage can be achieved by different

methods, described below. Most of the
methods contribute to crypsis, helping to
hide against a background; but mimesis
and motion dazzle protect without hiding.
Methods may be applied on their own or in
combination.

Crypsis

Crypsis means making the animal or

military equipment hard to see (or to
detect in other ways, such as by sound or
scent). Visual crypsis can be achieved in
many different ways, such as by living
underground or by being active only at
night, as well as by a variety of methods of
camouflage.[73]

Resemblance to the surroundings

Some animals' colours and patterns

resemble a particular natural background.
This is an important component of
camouflage in all environments. For
instance, tree-dwelling parakeets are
mainly green; woodcocks of the forest
floor are brown and speckled; reedbed
bitterns are streaked brown and buff; in
each case the animal's coloration matches
the hues of its habitat.[74][75] Similarly,
desert animals are almost all desert
coloured in tones of sand, buff, ochre, and
brownish grey, whether they are mammals
like the gerbil or fennec fox, birds such as
the desert lark or sandgrouse, or reptiles
like the skink or horned viper.[76]Military
uniforms, too, generally resemble their
backgrounds; for example khaki uniforms
are a muddy or dusty colour, originally
chosen for service in South
Asia.[77]Many[78] moths show industrial
melanism, including the peppered moth
which has coloration that blends in with
tree bark.[79] The coloration of these
insects evolved 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.[78][c] This is taken by zoologists as
evidence that camouflage is influenced by
natural selection, as well as demonstrating
that it changes where necessary to
resemble the local background.[78]
  

Black-faced sandgrouse is coloured like its

desert background.

Egyptian nightjar nests in open sand with

only its camouflaged plumage to protect
it.
  

Papuan frogmouth resembles a broken

branch.

Conspicuous giraffe mother can defend

herself, but calf hides for much of day,
relying on its camouflage.
  

Bright green katydid has the colour of

fresh vegetation.

Disruptive coloration

Illustration of the principle of "maximum disruptive

contrast" by Hugh 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,[80] or to
conceal telltale features, especially the
eyes, as in the common frog.[81] Disruptive
patterns may use more than one method
to defeat visual systems such as edge
detection.[82] Predators like the leopard
use disruptive camouflage to help them
approach prey, while potential prey like the
Egyptian nightjar use it to avoid detection
by predators.[83] 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.[84][85][86]

The presence of bold skin markings does

not in itself prove that an animal relies on
camouflage, as that depends on its
behaviour.[87] For example, although
giraffes have a high contrast pattern that
could be disruptive coloration, the adults
are extremely 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.[88] 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.[88]
A different explanation is implied by the
fact that young giraffes are far more
vulnerable to predation than adults: more
than half of all giraffe calves die within a
year,[88] and giraffe mothers hide their
calves, which spend much of the time lying
down in cover while their mothers are
away feeding. Since the presence of a
mother nearby does not affect survival, it
is argued that young giraffes must be
extremely well camouflaged; this is
supported by the fact that coat markings
are strongly inherited.[88]

Leopard: a disruptively camouflaged

predator

Russian T-90 battle tank painted in bold

disruptive pattern of sand and green
  

A ptarmigan and five chicks exhibit

exceptional disruptive camouflage

Jumping spider: a disruptively

camouflaged invertebrate predator

Eliminating shadow
 

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 the horned lizards

of North America, have evolved elaborate
measures to eliminate shadow. 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.[89] 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, the roundtail
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.[89] 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.[90] Similarly, some ground-nesting
birds including the European nightjar
select a resting position facing the sun.[90]
The elimination of shadow was identified
as a principle of military camouflage
during the Second World War.[91]
  

Three countershaded and cryptically

coloured ibex almost invisible in the Israeli
desert

"Shape, shine, shadow" make these

'camouflaged' military vehicles easily
visible.
  

The flat-tail horned lizard's body is

flattened and fringed to minimise its
shadow.

Camouflage netting is draped away from a

military vehicle to reduce its shadow.
  

A caterpillar's fringe of bristles conceals

its shadow.

Distraction

Conspicuous white distractive markings of bushbuck

(Tragelaphus scriptus take a predator's attention from
recognising the prey.
Many prey animals have conspicuous
high-contrast markings which
paradoxically attract the predator's
gaze.[d][92] These distractive markings
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 for
blue tits increased when artificial prey had
distractive markings.[93]

Self-decoration

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, a
caddis fly larva builds a decorated case
and lives almost entirely inside it; a
decorator crab covers its back with
seaweed, sponges and stones.[73] The
nymph of the predatory masked 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.[94][95]

Similar principles can be applied for

military purposes, for instance when a
sniper wears a ghillie 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.[96] Cott takes the example of the
larva of the blotched 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."[97][98]
  

This decorator crab has covered its body

with sponges.

Sniper in a Ghillie suit with plant materials

  

Crab camouflaged with algae

Reduvius personatus, masked hunter bug

nymph, camouflaged with sand grains
  

Soviet tanks under netting dressed with

vegetation, 1938

Cryptic behaviour

The leafy 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.[99] 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 the tiger, moving with
extreme stealth, both slowly and quietly,
watching its prey for any sign they are
aware of its presence.[99] 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.[88] The flat-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.[100] In
the ocean, the leafy sea dragon sways
mimetically, like the seaweeds amongst
which it rests, as if rippled by wind or
water currents.[101] 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.[102]

Motion camouflage

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.[103] However, insects such as
hoverflies[104] and dragonflies use motion
camouflage: the hoverflies to approach
possible mates, and the dragonflies to
approach rivals when defending
territories.[105][106] 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 to loom
larger in the target's field of vision.[107] The
same technique can be used for military
purposes, for example by missiles to
minimise their risk of detection by the
enemy.[104] However, missile engineers,
and animals such as bats, use the
technique primarily for its efficiency rather
than camouflage.[108]
  

Male Syritta pipiens hoverflies use motion

camouflage to approach females

Male Australian Emperor dragonflies use

motion camouflage to approach rivals.
 

Fish and frog melanophore cells change colour by

moving pigment-containing bodies.

Changeable skin coloration

Animals such as chameleon, frog,[109]

flatfish such as the peacock flounder,
squid and octopus actively change their
skin patterns and colours using special
chromatophore cells to resemble their
current background, or, as in most
chameleons, for signalling.[110] However,
Smith's dwarf chameleon does use active
colour change for camouflage.[111]

Four frames of the same peacock flounder taken a few

minutes apart, showing its ability to match its
coloration to the environment

Each chromatophore contains pigment of

only one colour. In fish and frogs, colour
change is mediated by the type of
chromatophores known as melanophores
that contain dark pigment. A melanophore
is star-shaped; it contains many small
pigmented organelles 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 by
hormones. In fish, the change is controlled
by the brain, which sends signals directly
to the chromatophores, as well as
producing hormones.[112]
The skins of cephalopods such as the
octopus contain complex units, each
consisting of a chromatophore with
surrounding muscle and nerve cells.[113]
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.[114][115]

On a longer timescale, animals like the

Arctic hare, Arctic fox, stoat, and rock
ptarmigan have snow 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 the dog family to do so.[116]
However, Arctic hares which live in the far
north of Canada, where summer is very
short, remain white year-round.[116][117]

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 techniques such as changing
uniforms and repainting vehicles for winter
have been in use since the Second World
War. In 2011, BAE Systems announced
their Adaptiv infrared camouflage
technology. It uses about 1000 hexagonal
panels to cover the sides of a tank. The
Peltier 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.[118][119][120]
  

Rock ptarmigan, changing colour in

springtime. The male is still mostly in
winter plumage

Norwegian volunteer soldiers in Winter

War, 1940, with white camouflage overalls
over their uniforms
  

Arctic hares in the low arctic change from

brown to white in winter

Snow-camouflaged German Marder III

jagdpanzer and white-overalled crew and
infantry in Russia, 1943
  

Veiled chameleon, Chamaeleo calyptratus,

changes colour primarily in relation to
mood and for signalling.

Adaptiv infrared camouflage lets an

armoured vehicle mimic a car.

Countershading
 

Countershading acts as a form of camouflage by

'painting out' the self-shadowing of the body or object.
The result is a 'flat' appearance, instead of the 'solid'
appearance 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.[121] Thayer observed that
"Animals are painted by Nature, darkest on
those parts which tend to be most lighted
by the sky's light, and vice versa".
Accordingly, the principle of
countershading is sometimes called
Thayer's Law.[122] Countershading is widely
used by terrestrial animals, such as
gazelles[123] and grasshoppers; marine
animals, such as sharks and dolphins;[124]
and birds, such as snipe and
dunlin.[125][126]

Countershading is less often used for

military camouflage, despite Second World
War experiments that showed its
effectiveness. English zoologist Hugh Cott
encouraged the use of techniques
including countershading, but despite his
authority on the subject, failed to persuade
the British authorities.[127] Soldiers often
wrongly viewed camouflage netting as a
kind of invisibility cloak, and they had to be
taught to look at camouflage practically,
from the enemy observer's point of
view.[128][129] At the same time in Australia,
zoologist William John Dakin advised
soldiers to copy animals' methods, using
their instincts for wartime camouflage.[130]
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 techniques used are the
matching of background colour and
pattern, and disruption of outlines.[123]
  

Countershaded Dorcas gazelle, Gazella

dorcas

Countershaded grey reef shark,

Carcharhinus amblyrhynchos
  

Countershaded ship and submarine in

Thayer's 1902 patent application

Two model birds painted by Thayer:

painted in background colours on the left,
countershaded and nearly invisible on the
right
  

Countershaded Focke-Wulf Fw 190D-9

Counter-illumination

Principle of counter-illumination in the firefly squid

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 of squid,
such as the firefly squid and the midwater
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.[131] Counterillumination camouflage
is the likely function of the
bioluminescence of many marine
organisms, though light is also produced
to attract[132] or to detect prey[133] and for
signalling.
Counterillumination has rarely been used
for military purposes. "Diffused lighting
camouflage" was trialled by Canada's
National 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.[134] The Canadian concept was
refined in the American Yehudi lights
project, and trialled in aircraft including B-
24 Liberators and naval Avengers.[135] The
planes were fitted with forward-pointing
lamps automatically adjusted to match the
brightness of the night sky.[134] This
enabled them to approach much closer to
a target – within 3,000 yards
(2,700 metres) – before being seen.[135]
Counterillumination was made obsolete by
radar, and neither diffused lighting
camouflage nor Yehudi lights entered
active service.[134]
  

HMS Largs by night with incomplete

diffused lighting camouflage, 1942, set to
maximum brightness

Bulwark of HMS Largs showing 4 (of about

60) diffused lighting fittings, 2 lifted, 2
deployed
  

Forward-looking Yehudi lights on

Grumman TBM Avenger torpedo bomber.

Yehudi Lights raise the average brightness

of the plane from a dark shape to the
same as the sky.

Transparency
 

Many animals of the open sea, like this Aurelia labiata

jellyfish, are largely transparent.

Many marine animals that float near the

surface are highly transparent, giving them
almost perfect camouflage.[136] However,
transparency is difficult for bodies made
of materials that have different refractive
indices from seawater. Some marine
animals such as jellyfish have gelatinous
bodies, composed mainly of water; their
thick mesogloea is acellular and highly
transparent. This conveniently makes
them buoyant, 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.[136] Gelatinous planktonic
animals are between 50 and 90 percent
transparent. A transparency of 50 percent
is enough to make an animal invisible to a
predator such as cod 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.[136]

Glass frogs like Hyalinobatrachium uranoscopum use

partial transparency for camouflage in the dim light of
the rainforest.
Some tissues such as muscles 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 vertebrate eye, which is made of the
protein crystallin, and the vertebrate
cornea which is made of the protein
collagen.[136] Other structures cannot be
made transparent, notably the retinas or
equivalent light-absorbing structures of
eyes — they must absorb light to be able to
function. The camera-type eye of
vertebrates and cephalopods must be
completely opaque.[136] Finally, some
structures are visible for a reason, such as
to lure prey. For example, the nematocysts
(stinging cells) of the transparent
siphonophore Agalma okenii resemble
small copepods.[136] Examples of
transparent marine animals include a wide
variety of larvae, including coelenterates,
siphonophores, salps (floating tunicates),
gastropod molluscs, polychaete worms,
many shrimplike crustaceans, and fish;
whereas the adults of most of these are
opaque and pigmented, resembling the
seabed or shores where they live.[136][137]
Adult comb 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 actually being
transparent.[137] The small Amazon river
fish Microphilypnus 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.[138]

Silvering
 

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 as sardine and herring are
camouflaged by silvering.[139]

The marine hatchetfish is extremely

flattened laterally, leaving the body just
millimetres thick, and the body is so silvery
as to resemble aluminium foil. The mirrors
consist of microscopic structures similar
to those used to provide structural
coloration: stacks of between 5 and 10
crystals of guanine spaced about ¼ of a
wavelength apart to interfere
constructively and achieve nearly 100 per
cent reflection. In the deep waters that the
hatchetfish lives in, only blue light with a
wavelength of 500 nanometres percolates
down and needs to be reflected, so mirrors
125 nanometres apart provide good
camouflage.[139]

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.[139] Silvering is found in
other marine animals as well as fish. The
cephalopods, including squid, octopus and
cuttlefish, have multi-layer mirrors made of
protein rather than guanine.[139]

Mimesis

In mimesis (also called masquerade), the

camouflaged object looks like something
else which is of no special interest to the
observer.[140] Mimesis is common in prey
animals, for example when a peppered
moth caterpillar mimics a twig, or a
grasshopper mimics a dry leaf.[141] It is
also found in nest structures; some
eusocial wasps, such as Leipomeles
dorsata, build a nest envelope in patterns
that mimic the leaves surrounding the
nest.[142]

Mimesis is also employed by some

predators and parasites to lure their prey.
For example, a flower mantis mimics a
particular kind of flower, such as an
orchid.[143] This tactic has occasionally
been used in warfare, for example with
heavily armed Q-ships disguised as
merchant ships.[144][145][146]

The common cuckoo, a brood 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 a
sparrowhawk. 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.[147]
The cuckoo's egg itself mimics the eggs of
the host species, reducing its chance of
being rejected.[148][149]
  

Peppered moth caterpillars mimic twigs

Flower mantis lures its insect prey by

mimicking a Phalaenopsis orchid blossom
  

Hooded grasshopper Teratodus

monticollis, superbly mimics a leaf with a
bright orange border

This grasshopper hides from predators by

mimicking a dry leaf
  

WWII tank hid from the enemy in Operation

Bertram by mimicking a truck

Armed WW1 Q-ship lured enemy

submarines by mimicking a merchantman
  

Cuckoo adult mimics sparrowhawk, giving

female time to lay eggs parasitically

Cuckoo eggs mimicking smaller eggs, in

this case of reed warbler
  

Wrap-around spider Dolophones

mimicking a stick

Motion dazzle

The zebra'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.[150] Motion dazzle may
degrade predators' ability to estimate the
prey's speed and direction accurately,
giving the prey an improved chance of
escape.[151] 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.[150] 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.[151]

An animal that is commonly thought to be

dazzle-patterned is the zebra. The bold
stripes of the zebra have been claimed to
be disruptive camouflage,[152] background-
blending and countershading.[153][e] After
many years in which the purpose of the
coloration was disputed,[154] an
experimental study by Tim Caro suggested
in 2012 that the pattern reduces the
attractiveness of stationary models to
biting flies such as horseflies and tsetse
flies.[155][156] 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 two visual illusions: the wagon-wheel
effect, where the perceived motion is
inverted, and the barberpole illusion, where
the perceived motion is in a wrong
direction.[157]

Civil applications
 

Cellphone tower disguised as a tree

A hide used in field sports

Camouflage is occasionally used to make

built structures less conspicuous: for
example, in South 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 actually on other structures.[158]

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.[159] Field
sports such as driven grouse shooting
conceal hunters in hides (also called
blinds or shooting butts).[160] 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.[161]

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.[162]

Fashion, art and society

The "dazzle ball" held by the Chelsea Arts Club, 1919

Military camouflage patterns influenced

fashion and art from the time of the First
World War onwards. Gertrude Stein
recalled the cubist artist Pablo 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.
 — Gertrude Stein in From Picasso
(1938)[163]

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.[164] The
Illustrated London News
announced:[164][165]

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.[166] 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.[167]

Modern artists such as Ian Hamilton Finlay

have used camouflage to reflect on war.
His 1973 screenprint of a tank
camouflaged in a leaf pattern, Arcadia, is
described by the Tate as drawing "an ironic
parallel between this idea of a natural
paradise and the camouflage patterns on a
tank".[168] The title refers to the Utopian
Arcadia of poetry and art, and the
memento mori Latin phrase Et in Arcadia
ego which recurs in Hamilton Finlay's
work. In science fiction, Camouflage is a
novel about shapeshifting alien beings by
Joe Haldeman.[169] 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.[170]
  

André Mare's Cubist sketch, c. 1917, of a

280 calibre gun illustrates the interplay of
art and war, as artists like Mare
contributed their skills as wartime
camoufleurs.

Ian Hamilton Finlay's 1973 Arcadia

screenprint uses camouflage in modern
art to contrast leafy peace and military
 hardware.

Camouflage clothing in an anti-war

protest, 1971

A camouflage patterned skirt as a fashion

item
Notes
a. A letter from Alfred Russel Wallace to
Darwin of March 8, 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]
b. Cott explains Beddard's observation as a
coincident disruptive pattern.[9]
c. Before 1860, unpolluted tree trunks were
often covered in pale lichens; polluted
trunks were bare, and often nearly black.
d. These distraction markings are
sometimes called dazzle markings, but
have nothing to do with motion dazzle or
wartime dazzle painting.
e. 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 primary
function of the stripes is camouflage.

References
1. Aristotle (c. 350 BC). Historia Animalium.
IX, 622a: 2–10. Cited in Borrelli, Luciana;
Gherardi, Francesca; Fiorito, Graziano
(2006). A catalogue of body patterning in
Cephalopoda. Firenze University Press.
ISBN 978-88-8453-377-7. Abstract
2. Darwin 1859.
3. Darwin 1859, p. 84.
4. Poulton 1890, p. 111.
5. Alfred Russel Wallace (8 March 1868).
"Alfred Russel Wallace Letters and
Reminiscences By James Marchant" .
Darwin Online. Retrieved 29 March 2013.
6. Poulton 1890, p. Fold-out after p. 339.
7. Beddard 1892, p. 83.
8. Beddard 1892, p. 87.
9. Cott 1940, pp. 74–75.
10. Beddard 1892, p. 122.
11. Thayer 1909.
12. Forbes 2009, p. 77.
13. Thayer 1909, pp. 5, 16.
14. Rothenberg 2011, pp. 132–133.
15. Wright, Patrick (23 June 2005). "Cubist
Slugs. Review of DPM: Disruptive Pattern
Material; An Encyclopedia of Camouflage:
Nature – Military – Culture by Roy
Behrens" . London Review of Books. 27
(12): 16–20.
16. Cott 1940, pp. 172–173.
17. Cott 1940, pp. 47–67.
18. Cott 1940, pp. 174–186.
19. Forbes 2009, pp. 153–155.
20. Lindgren, Johan and Peter Sjövall, Ryan
M. Carney, Per Uvdal, Johan A. Gren, Gareth
Dyke, Bo Pagh Schultz, Matthew D.
Shawkey, Kenneth R. Barnes, Michael J.
Polcyn (February 2014). "Skin pigmentation
provides evidence of convergent melanism
in extinct marine reptiles". Nature. 506
(7489): 484–488.
doi:10.1038/nature12899 .
PMID 24402224 .
21. Casson 1995, pp. 211–212.
22. Casson 1995, p. 235.
23. 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 . JSTOR 2563673 .
24. Payne-Gallwey, Ralph (1903). The
Crossbow. Longmans, Green. p. 11.
25. Saunders, Nicholas (2005). The People
of the Caribbean: An Encyclopedia of
Archaeology and Traditional Culture. ABC-
CLIO.
26. Haythornthwaite, P (2002). British
Rifleman 1797–1815. Osprey Publishing.
p. 20. ISBN 978-1841761770.
27. Newark 2007, p. 43.
28. "Killers in Green Coats" . Weider History
Group. 20 February 2008. Retrieved 8 July
2012.
29. "Khaki Uniform 1848-49: First
Introduction by Lumsden and Hodson".
Journal of the Society for Army Historical
Research. 82 (Winter): 341–347. 2004.
30. Hodson, W.S.R.; edited by George H.
Hodson (1859). Twelve Years of a Soldier's
Life in India, being extracts from the letters
of the late Major WSR Hodson . John W.
Parker and Son.
31. Barthorp, Michael (1988). The British
Army on Campaign 1816–1902. 4 (1882–
1902). Osprey Publishing. pp. 24–33.
ISBN 0-85045-849-8.
32. Chappell, M (2003). The British Army in
World War I (1). Osprey Publishing. p. 37.
ISBN 978-1-84176-399-6.
33. Wright, Patrick (23 June 2005). "Cubist
Slugs" . London Review of Books. 27 (12):
16–20.
34. Guirand de Scévola, Lucien-Victor
(December 1949). "Souvenir de
Camouflage (1914–1918)". Revue des Deux
Mondes (in French).
35. Forbes, 2009. pp. 104–105.
36. "Art of the First World War: André Mare
and Leon Underwood" . The Elm at
Vermezeele. Memorial-Caen. 1998.
Retrieved 8 February 2013.
37. "Art of the First World War: André
Mare" . Memorial-Caen. 1998. Retrieved
8 February 2013.
38. "Camouflage" . Online Etymology
Dictionary. 2012. Retrieved 8 February
2013.
39. "Camouflage, n" . Oxford English
Dictionary. Oxford University Press.
Retrieved 8 February 2013.
40. Forbes 2009, pp. 85–89.
41. For Solomon, see BBC Radio 4
programme "Warpaint: the story of
camouflage" by Patrick Wright, August
2002 (repeated Radio 4 Extra, 17 June
2014).
42. Sumrall, RF (February 1973). "Ship
Camouflage (WWII): Deceptive Art". United
States Naval Institute Proceedings: 67–81.
43. Prinzeugen. "Schnellboot: An Illustrated
Technical History" . Prinz Eugen. Retrieved
5 March 2012.
44. "Obituary: Mr Norman Wilkinson,
Inventor of 'dazzle' painting". The Times. 1
June 1971. p. 12.
45. Wilkinson, Norman (1969). A Brush with
Life. Seeley Service. p. 79.
46. Forbes 2009, pp. 149–150.
47. Keating, Kenneth C (1981).
"Maskirovka: The Soviet System of
Camouflage" (PDF). U.S. Army Russian
Institute. Retrieved 8 July 2012.
48. Clark, Lloyd (2011). Kursk: the greatest
battle. Headline Review. p. 278. ISBN 978-0-
7553-3639-5.
49. Shaw, Robert L. (1985). Fighter
Combat : Tactics and Maneuvering. Naval
Institute Press. ISBN 0-87021-059-9.
50. 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.
51. Tinbergen, Niko (1953). The Herring
Gull's World. Collins. p. 14. ISBN 0-00-
219444-9. “white has proved to be the most
efficient concealing coloration for aircraft
on anti-submarine patrol”
52. "World War II" . Farnham Castle.
Retrieved 8 February 2013.
53. Forbes 2009, pp. 151–152.
54. Barkas 1952, pp. 154, 186–188.
55. Forbes 2009, pp. 156–166.
56. Mellor, D P, , Australia in the War of
1939–1945. Series 4 – Civil, Volume 5 The
Role of Science and Industry, Australian
War Memorial, Canberra, 1968, page 538ff
57. "Concealment, Camouflage, and
Deception" (PDF). Smithsonian. pp. 1–4.
Retrieved 16 June 2012.
58. "FM 21–75" . Chapter 1: Cover,
Concealment, and Camouflage.
Department of the Army. Retrieved 16 June
2012.
59. "FM 21-305/AFMAN 24-306" (PDF).
Chapter 20: Vehicle Camouflage And
Nuclear, Biological, And Chemical
Operations. Department of the Army. pp. 1–
9. Retrieved 16 June 2012.
60. "5–103" . Appendix D: Camouflage.
Department of the Army. Retrieved 17 June
2012.
61. "SSZ Camouflage" . Military Suppliers &
News. 2012. Retrieved 17 June 2012.
62. Jukkola, EE; Cohen, R. (1946). "Color
Stability of Olive Drab Infrared-Reflecting
Camouflage Finishes" . Industrial &
Engineering Chemistry. 38 (9): 927–930.
doi:10.1021/ie50441a019 .
63. Richardson, Doug (2001). Stealth
Warplanes: Deception, Evasion, and
Concealment in the Air . MBI Publishing /
Zenith Press. ISBN 978-0-7603-1051-9.
64. Pfanner, Toni (March 2004). "Military
uniforms and the law of war" (PDF). IRRC.
86 (853): 99–100.
doi:10.1017/s1560775500180113 .
65. FM 21–76 US Army Survival Manual .
Department of the Army. Retrieved
8 January 2013.
66. Photosimulation Camouflage Detection
Test . U.S. Army Natick Soldier Research,
Development and Engineering Center. 2009.
p. 27. Retrieved 5 October 2012.
67. Brayley, Martin J (2009). Camouflage
uniforms: international combat dress
1940–2010. Crowood. ISBN 1-84797-137-7.
68. Freedberg, SJ Jr. (25 June 2012). "Army
drops universal camouflage after spending
billions" . AOL Defence. Archived from the
original on 31 August 2012. Retrieved
27 September 2012.
69. Davies, W. "Berlin Brigade Urban Paint
Scheme" . Newsletter. Ex-Military Land
Rover Association. Retrieved 25 September
2012.
70. Craemer, Guy. "Dual Texture – U.S.
Army digital camouflage" . United
Dynamics. Retrieved 27 September 2012.
71. Gye, H (25 June 2012). "How U.S. Army
spent $5 billion on 'failed' pixel
camouflage" . Daily Mail. Retrieved
21 November 2012.
72. Engber, D. (5 July 2012) [2007]. "Lost in
the Wilderness, the military's
misadventures in pixellated camouflage" .
Slate. Retrieved 27 September 2012.
73. Forbes 2009, pp. 50–51 and passim.
74. Cott 1940, pp. 5–19.
75. Forbes 2009, p. 51.
76. Cott 1940, pp. 5–6.
77. Newark 2007, pp. 45–46.
78. Cott 1940, p. 17.
79. Still, J (1996). Collins Wild Guide:
Butterflies and Moths. HarperCollins.
p. 158. ISBN 0-00-220010-4.
80. Barbosa, A; Mathger, LM; Buresch, KC;
Kelly, J; Chubb, C; Chiao, C; Hanlon RT
(2008). "Cuttlefish camouflage: The effects
of substrate contrast and size in evoking
uniform, mottle or disruptive body
patterns". Vision Research. 48 (10): 1242–
1253. doi:10.1016/j.visres.2008.02.011 .
PMID 18395241 .
81. Cott 1940, pp. 83–91.
82. Osorio, Daniel; Cuthill, Innes C.
"Camouflage and perceptual organization
in the animal kingdom" (PDF). Retrieved
25 October 2013.
83. Stevens, M; Cuthill, IC; Windsor, AMM;
Walker, HJ (7 October 2006). "Disruptive
contrast in animal camouflage" .
Proceedings of the Royal Society B. 273
(1600): 2433–2436.
doi:10.1098/rspb.2006.3614 .
PMC 1634902  . PMID 16959632 .
84. Sweet, KM (2006). Transportation and
Cargo Security: Threats and Solutions.
Prentice Hall. p. 219. ISBN 978-0-13-
170356-8.
85. FM 5–20: Camouflage, Basic
Principles . U.S. War Department.
November 2015 [1944].
86. Field Manual Headquarters No. 20-3 .
Camouflage, Concealment, and Decoys.
Department of the Army. 30 August 1999.
87. Roosevelt, Theodore (1911). "Revealing
and concealing coloration in birds and
mammals" . Bulletin of the American
Museum of Natural History. 30 (Article 8):
119–231. Roosevelt attacks Thayer on
page 191, arguing that neither zebra nor
giraffe are "'adequately obliterated' by
countershading or coloration pattern or
anything else."
88. Mitchell, G; Skinner, JD (2003). "On the
origin, evolution and phylogeny of giraffes
Giraffa camelopardalis" (PDF).
Transactions of the Royal Society of South
Africa. 58 (1): 51–73.
doi:10.1080/00359190309519935 .
89. Sherbrooke, WC (2003). Introduction to
horned lizards of North America .
University of California Press. pp. 117–118.
ISBN 978-0-520-22825-2.
90. Cott 1940, pp. 104–105.
91. U.S. War Department (November 1943).
"Principles of Camouflage" . Tactical and
Technical Trends (37).
92. Stevens, M.; Merilaita, S. (2009).
"Defining disruptive coloration and
distinguishing its functions". Philosophical
Transactions of the Royal Society B:
Biological Sciences. 364 (1516): 481–488.
doi:10.1098/rstb.2008.0216 .
93. Dimitrova, M.; Stobbe, N.; Schaefer, H.
M.; Merilaita, S. (2009). "Concealed by
conspicuousness: distractive prey
markings and backgrounds". Proceedings
of the Royal Society B: Biological Sciences.
276 (1663): 1905–1910.
doi:10.1098/rspb.2009.0052 .
94. Wierauch, C. (2006). "Anatomy of
disguise: camouflaging structures in
nymphs of Some Reduviidae
(Heteroptera)". American Museum
Novitates. 3542: 1–18. doi:10.1206/0003-
0082(2006)3542[1:AODCSI]2.0.CO;2 .
95. Bates, Mary. "Natural Bling: 6 Amazing
Animals That Decorate Themselves" .
National Geographic. Retrieved 11 June
2015.
96. Forbes 2009, pp. 102–103.
97. Cott 1940, p. 360.
98. Ruxton, Graeme D.; Stevens, Martin (1
June 2015). "The evolutionary ecology of
decorating behaviour" . Biology Letters. 11
(6): 20150325.
doi:10.1098/rsbl.2015.0325 .
99. Cott 1940, p. 141.
100. "What is a Horned Lizard?" .
hornedlizards.org. Horned Lizard
Conservation Society. Retrieved
14 November 2015.
101. "Leafy Sea Dragon" . WWF. Retrieved
21 December 2011.
102. Bian, Xue; Elgar, Mark A.; Peters,
Richard A. (2016). "The swaying behavior of
Extatosoma tiaratum : motion camouflage
in a stick insect?" . Behavioral Ecology. 27
(1): 83–92. doi:10.1093/beheco/arv125 .
103. Cott 1940, pp. 141–143.
104. Srinivasan, MV; Davey, M (1995).
"Strategies for active camouflage of
motion". Proceedings of the Royal Society
B. 259 (1354): 19–25.
doi:10.1098/rspb.1995.0004 .
105. Hopkin, Michael (5 June 2003).
"Dragonfly flight tricks the eye" .
Nature.com. Retrieved 16 January 2012.
106. Mizutani, AK; Chahl, JS; Srinivasan,
MV (5 June 2003). "Insect behaviour:
Motion camouflage in dragonflies" . Nature.
65 (423): 604. doi:10.1038/423604a .
107. Glendinning, P (2004). "The
mathematics of motion camouflage" .
Proceedings of the Royal Society B. 271
(1538): 477–481.
doi:10.1098/rspb.2003.2622 .
PMC 1691618  . PMID 15129957 .
108. Ghose, K; Horiuchi, TK; Krishnaprasad,
PS; Moss, CF (2006). "Echolocating Bats
Use a Nearly Time-Optimal Strategy to
Intercept Prey" . PLoS Biology. 4 (5): e108.
doi:10.1371/journal.pbio.0040108 .
PMC 1436025  . PMID 16605303 .
109. Cott 1940, pp. 30–31.
110. Forbes 2009, pp. 52, 236.
111. Stuart-Fox, Devi; Moussalli, Adnan;
Whiting, Martin J. (23 August 2008).
"Predator-specific camouflage in
chameleons" . Biology Letters. 4 (4).
doi:10.1098/rsbl.2008.0173 .
112. Wallin, M (2002). "Nature's Palette"
(PDF). Bioscience Explained. 1 (2): 1–12.
Retrieved 17 November 2011.
113. Cott, 1940. p. 32.
114. Cloney, RA; Florey, E (1968).
"Ultrastructure of Cephalopod
Chromatophore Organs". Zeitschrift für
Zellforschung und mikroskopische
Anatomie. 89 (2): 250–80.
doi:10.1007/BF00347297 .
PMID 5700268 .
115. "Day Octopuses, Octopus cyanea" .
MarineBio Conservation Society. Retrieved
31 January 2013.
116. "Arctic Wildlife" . Churchill Polar Bears.
2011. Retrieved 22 December 2011.
117. Hearn, Brian (20 February 2012). The
Status of Arctic Hare (Lepus arcticus
bangsii) in Insular Newfoundland (PDF).
Newfoundland Labrador Department of
Environment and Conservation. p. 7.
Retrieved 3 February 2013.
118. "Tanks test infrared invisibility cloak" .
BBC News. 5 September 2011. Retrieved
13 June 2012.
119. "Adaptiv – A Cloak of Invisibility" . BAE
Systems. 2011. Retrieved 14 November
2015.
120. "Innovation Adaptiv Car Signature" .
BAE Systems. 2012. Retrieved
14 November 2015.
121. Cott 1940, pp. 35–46.
122. Forbes 2009, pp. 72–73.
123. Kiltie, 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 .
124. Cott 1940, p. 41.
125. Ehrlich, Paul R; Dobkin, David S;
Wheye, Darryl (1988). "The Color of Birds" .
Stanford University. Retrieved 1 February
2013.
126. Cott 1940, p. 40.
127. Forbes 2009, pp. 146–150.
128. Forbes 2009, p. 152.
129. Barkas 1952, p. 36.
130. Elias 2011, pp. 57–66.
131. "Midwater Squid, Abralia veranyi" .
Smithsonian National Museum of Natural
History. Retrieved 28 November 2011.
132. Young, Richard Edward (October
1983). "Oceanic Bioluminescence: an
Overview of General Functions" . Bulletin of
Marine Science. 33 (4): 829–845.
133. Douglas, RH; Mullineaux, CW;
Partridge, JC (September 2000). "Long-
wave sensitivity in deep-sea stomiid
dragonfish with far-red bioluminescence:
evidence for a dietary origin of the
chlorophyll-derived retinal photosensitizer
of Malacosteus niger" . Philosophical
Transactions of the Royal Society B. 355
(1401): 1269–1272.
doi:10.1098/rstb.2000.0681 .
PMC 1692851  . PMID 11079412 .
134. "Diffused Lighting and its use in the
Chaleur Bay" . Naval Museum of Quebec.
Royal Canadian Navy. Archived from the
original on 22 May 2013. Retrieved
3 February 2013.
135. Hambling, David (9 May 2008). "Cloak
of Light Makes Drone Invisible?" . Wired.
Retrieved 17 June 2012.
136. Herring 2002, pp. 190–191.
137. Cott 1940, p. 6.
138. Carvalho, Lucélia Nobre; Zuanon,
Jansen; Sazima, Ivan (April–June 2006).
"The almost invisible league: crypsis and
association between minute fishes and
shrimps as a possible defence against
visually hunting predators" . Neotropical
Ichthyology. 4 (2): 219–224.
doi:10.1590/S1679-62252006000200008 .
139. Herring 2002, pp. 192–195.
140. Gullan, PJ; Cranston, PS (2010). The
Insects (4th ed.). John Wiley, Blackwell.
pp. 512–513. ISBN 978-1-4443-3036-6.
141. Forbes 2009, p. 151.
142. Ross, Kenneth G. (1991). The Social
Biology of Wasps. Cornell Press. p. 233.
ISBN 978-0-801-49906-7.
143. Forbes 2009, p. 134.
144. Beyer, Kenneth M (1999). Q-Ships
versus U-Boats: America's Secret Project.
Naval Institute Press. ISBN 1-55750-044-4.
145. McMullen, Chris (2001). "Royal Navy
'Q' Ships" . Great War Primary Documents
Archive. Retrieved 6 March 2012.
146. Forbes 2009, pp. 6–42.
147. Welbergen, J; Davies, NB (2011). "A
parasite in wolf's clothing: hawk mimicry
reduces mobbing of cuckoos by hosts".
Behavioral Ecology. 22 (3): 574–579.
doi:10.1093/beheco/arr008 .
148. Brennand, Emma (24 March 2011).
"Cuckoo in egg pattern 'arms race' " . BBC
News. Retrieved 22 August 2011.
149. 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 .
150. Scott-Samuel, NE; Baddeley, R; Palmer,
CE; Cuthill, IC (June 2011). Burr, David C,
ed. "Dazzle Camouflage Affects Speed
Perception" . PLOS ONE. 6 (6): e20233.
doi:10.1371/journal.pone.0020233 .
PMC 3105982  . PMID 21673797 .
151. Stevens, M; Searle, WTL; Seymour, JE;
Marshall, KLA; Ruxton, GD (25 November
2011). "Motion dazzle and camouflage as
distinct anti-predator defenses" . BMC
Biology. pp. 9–81. doi:10.1186/1741-7007-
9-81 .
152. Cott 1940, p. 94.
153. Thayer 1909, p. 136.
154. Caro, Tim (2009). "Contrasting
coloration in terrestrial mammals" .
Philosophical Transactions of the Royal
Society B. 364 (1516): 537–548.
doi:10.1098/rstb.2008.0221 .
PMC 2674080  . PMID 18990666 .
155. 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.
156. Egri, Ádám; Miklós Blahó; György
Kriska; Róbert Farkas; Mónika Gyurkovszky;
Susanne Åkesson; Gábor Horváth (March
2012). "Polarotactic tabanids find striped
patterns with brightness and/or
polarization modulation least attractive: an
advantage of zebra stripes" . The Journal
of Experimental Biology. 215 (5): 736–745.
doi:10.1242/jeb.065540 . PMID 22323196 .
157. How, Martin J.; Zanker, Johannes M.
(2014). "Motion camouflage induced by
zebra stripes" . Zoology. 117: TBA.
doi:10.1016/j.zool.2013.10.004 .
158. du Plessis, A (3 July 2002).
Telecommunication Mast Management
Guidelines For The City of Tshwane . City of
Tshwane Metropolitan Municipality.
159. Newark 2007, p. 38.
160. Blakeley, Peter F (2012).
Wingshooting. Stackpole Books. pp. 116,
125. ISBN 978-0-8117-0566-0.
161. Newark 2007, pp. 48, 50.
162. "The secrets behind all that
camouflage" . Automotive News. 12 May
2015. Retrieved 28 July 2015.
163. Stein, Gertrude; Toklas, Alice B.
(trans.) (1939). "Picasso" . Scribners.
164. Forbes 2009, p. 100.
165. "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.
166. "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 from the original
on 12 December 2012. Retrieved
1 December 2011.
167. "Camouflage: The Exhibition" .
Canadian War Museum. 5 June 2009.
Retrieved 14 November 2015.
168. "Ian Hamilton Finlay: Arcadia
(collaboration with George Oliver)" .
Arcadia, 1973. Tate. July 2008. Retrieved
11 May 2012.
169. Haldeman, Joe (2004). Camouflage.
Ace Books. ISBN 0-441-01161-6.
170. Frank, Thaisa (1992). A Brief History
of Camouflage. Black Sparrow Press.
ISBN 0-87685-857-4.
Bibliography
Camouflage in nature

Early research
Beddard, Frank Evers (1892). Animal
Coloration. Swan Sonnenschein.
Cott, Hugh B. (1940). Adaptive
Coloration in Animals. Methuen.
Darwin, Charles (1859). On the Origin of
Species. John Murray. Reprinted 1985,
Penguin Classics.
Poulton, Edward B. (1890). The Colours
of Animals. Kegan Paul, Trench, Trübner.
Thayer, Abbott Handerson (1909).
Concealing-Coloration in the Animal
Kingdom Macmillan.
General reading
Elias, Ann (2011). Camouflage Australia:
Art, Nature, Science and War Sydney
University Press. ISBN 978-1-920899-73-
8.
 Forbes, Peter (2009). Dazzled and
Deceived: Mimicry and Camouflage Yale
University Press. ISBN 978-0-300-17896-
8.
Herring, Peter (2002). The Biology of the
Deep Ocean Oxford University Press.
ISBN 978-0-19-854956-7.
Rothenberg, David (2011). Survival of the
Beautiful: Art, Science and Evolution
Bloomsbury. ISBN 978-1-60819-216-8.

Military camouflage

Barkas, Geoffrey (1952). The

Camouflage Story (from Aintree to
Alamein). Cassell.
 Casson, Lionel (1995). Ships and
Seamanship in the Ancient World. JHU
Press. ISBN 978-0-8018-5130-8.
Newark, Tim (2007). Camouflage.
Thames and Hudson, with Imperial War
Museum. ISBN 978-0-500-51347-7.

Further reading
Behrens, Roy R. (2002). False Colors: Art,
Design and Modern Camouflage.
Bobolink Books. ISBN 0-9713244-0-9.
Behrens, Roy R. (2009). Camoupedia: A
Compendium of Research on Art,
Architecture and Camouflage. Bobolink
Books. ISBN 978-0-9713244-6-6.
Behrens, Roy R. (editor) (2012). Ship
Shape: A Dazzle Camouflage
Sourcebook. Bobolink Books. ISBN 978-
0-9713244-7-3.
Goodden, Henrietta (2009). Camouflage
and Art: Design for Deception in World
War 2. Unicorn Press. ISBN 978-0-
906290-87-3.
Latimer, Jon (2001). Deception in War.
John Murray. ISBN 978-1-58567-381-0.
Newman, Alex; Blechman, Hardy (2004).
DPM – Disruptive Pattern Material: An
Encyclopaedia of Camouflage: Nature,
Military and Culture. DPM. ISBN 978-0-
9543404-0-7.
Stevens, Martin; Merilaita, Sami (2011).
Animal Camouflage: Mechanisms and
Function. Cambridge University Press.
ISBN 978-0-521-15257-0.
Wickler, Wolfgang (1968). Mimicry in
plants and animals. McGraw-Hill.
ISBN 978-0-07-070100-7.
For children
Kalman, Bobbie; Crossingham, John
(2001). What are Camouflage and
Mimicry?. Crabtree Publishing.
ISBN 978-0-86505-962-7. (ages 4–8)
Mettler, Rene (2001). Animal
Camouflage. First Discovery series.
 Moonlight Publishing. ISBN 978-1-
85103-298-3. (ages 4–8)

External links
Wikimedia Commons has media related
 
to Camouflage.

Wikimedia Commons has media related

 
to Animal camouflage.

Wikisource has the text of the 1922

  Encyclopædia Britannica article
Camouflage.

Ohio State University: The Camouflage

Project – interplay of science and art
 Behrens, Roy. A Chronology of
Camouflage
"An informal study into camouflage"

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