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Motion camouflage is
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 ...
which provides a degree of concealment for a moving object, given that motion makes objects easy to detect however well their coloration matches their background or breaks up their outlines. The principal form of motion camouflage, and the type generally meant by the term, involves an attacker's mimicking the
optic flow Optical flow or optic flow is the pattern of apparent motion of objects, surfaces, and edges in a visual scene caused by the relative motion between an observer and a scene. Optical flow can also be defined as the distribution of apparent veloci ...
of the background as seen by its target. This enables the attacker to approach the target while appearing to remain stationary from the target's perspective, unlike in classical pursuit (where the attacker moves straight towards the target at all times, and often appears to the target to move sideways). The attacker chooses its flight path so as to remain on the line between the target and some landmark point. The target therefore does not see the attacker move from the landmark point. The only visible evidence that the attacker is moving is its
looming ''Looming'' is a term found in the study of perception, as it relates directly to psychology. Looming occurs when an object begins moving closer to the eye. As the resulting image becomes increasingly larger on the perceiver's retina, i.e., when ...
, the change in size as the attacker approaches. Motion is also used in a variety of other camouflage strategies, including swaying to mimic plant movements in the wind or ocean currents. First discovered in
hoverflies Hover flies, also called flower flies or syrphid flies, make up the insect family Syrphidae. As their common name suggests, they are often seen hovering or nectaring at flowers; the adults of many species feed mainly on nectar and pollen, while ...
in 1995, motion camouflage by minimising optic flow has been demonstrated in another insect order,
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threate ...
, as well as in two groups of
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
,
falcon Falcons () are birds of prey in the genus ''Falco'', which includes about 40 species. Falcons are widely distributed on all continents of the world except Antarctica, though closely related raptors did occur there in the Eocene. Adult falcons ...
s and echolocating
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
s. Since bats hunting at night cannot be using the strategy for camouflage, it has been named, describing its mechanism, as constant absolute target direction. This is an efficient homing strategy, and it has been suggested that anti-aircraft missiles could benefit from similar techniques. Camouflage is sometimes facilitated by motion, as in the
leafy sea dragon The leafy seadragon (''Phycodurus eques'') or Glauert's seadragon, is the only member of the genus ''Phycodurus'' and is a marine fish in the family Syngnathidae, which includes seadragons, pipefish, and seahorses. It is found along the sou ...
and some
stick insect The Phasmatodea (also known as Phasmida, Phasmatoptera or Spectra) are an order of insects whose members are variously known as stick insects, stick-bugs, walking sticks, stick animals, or bug sticks. They are also occasionally referred to as ...
s. These animals complement their passive camouflage by swaying like plants, delaying their recognition by predators.


Camouflage of approach motion

Many animals are highly sensitive to motion; for example,
frog A frog is any member of a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-frog" ''Triadobatrachus'' is ...
s readily detect small moving dark spots but ignore stationary ones. Therefore, motion signals can be used to defeat camouflage. Moving objects with
disruptive camouflage Disruptive coloration (also known as disruptive camouflage or disruptive patterning) is a form of camouflage that works by breaking up the outlines of an animal, soldier or military vehicle with a strongly contrasting pattern. It is often comb ...
patterns remain harder to identify than uncamouflaged objects, especially if other similar objects are nearby, even though they are detected, so motion does not completely 'break' camouflage. All the same, the conspicuousness of motion raises the question of whether and how motion itself could be camouflaged. Several mechanisms are possible.


Stealthy movements

One strategy is to minimise actual motion, as when
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
s such as
tiger The tiger (''Panthera tigris'') is the largest living cat species and a member of the genus ''Panthera''. It is most recognisable for its dark vertical stripes on orange fur with a white underside. An apex predator, it primarily preys on un ...
s stalk prey by moving very slowly and stealthily. This strategy effectively avoids the need to camouflage motion.


Minimising motion signal

When movement is required, one strategy is to minimise the motion signal, for example by avoiding waving limbs about and by choosing patterns that do not cause flicker when seen by the prey from straight ahead.
Cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control o ...
may be doing this with their
active camouflage Active camouflage or adaptive camouflage is camouflage that adapts, often rapidly, to the surroundings of an object such as an animal or military vehicle. In theory, active camouflage could provide perfect concealment from visual detection. Activ ...
by choosing to form stripes at right angles to their front-back axis, minimising motion signals that would be given by occluding and displaying the pattern as they swim.


Disrupting perception of motion

Disrupting the attacker's perception of the target's motion was the main intended purpose of
dazzle camouflage Dazzle camouflage, also known as razzle dazzle (in the U.S.) or dazzle painting, is a family of ship camouflage that was used extensively in World War I, and to a lesser extent in World War II and afterwards. Credited to the British marine a ...
as used on ships in the
First World War World War I (28 July 1914 11 November 1918), often abbreviated as WWI, was one of the deadliest global conflicts in history. Belligerents included much of Europe, the Russian Empire, the United States, and the Ottoman Empire, with fighti ...
, though its effectiveness is disputed. This type of dazzle does not appear to be used by animals.


Mimicking optic flow of background

Some animals mimic the
optic flow Optical flow or optic flow is the pattern of apparent motion of objects, surfaces, and edges in a visual scene caused by the relative motion between an observer and a scene. Optical flow can also be defined as the distribution of apparent veloci ...
of the background, so that the attacker does not appear to move when seen by the target. This is the main focus of work on motion camouflage, and is often treated as synonymous with it.


Pursuit strategies

An attacker can mimic the background's optic flow by choosing its flight path so as to remain on the line between the target and either some real landmark point, or a point at infinite distance (giving different pursuit algorithms). It therefore does not move from the landmark point as seen by the target, though it inevitably looms larger as it approaches. This is not the same as moving straight towards the target (classical pursuit): that results in visible sideways motion with a readily detectable difference in optic flow from the background. The strategy works whether the background is plain or textured. This motion camouflage strategy was discovered and modelled as
algorithm In mathematics and computer science, an algorithm () is a finite sequence of rigorous instructions, typically used to solve a class of specific problems or to perform a computation. Algorithms are used as specifications for performing ...
s in 1995 by M. V. Srinivasan and M. Davey while they were studying mating behaviour in
hoverflies Hover flies, also called flower flies or syrphid flies, make up the insect family Syrphidae. As their common name suggests, they are often seen hovering or nectaring at flowers; the adults of many species feed mainly on nectar and pollen, while ...
. The male hoverfly appeared to be using the tracking technique to approach prospective mates. Motion camouflage has been observed in high-speed territorial battles between
dragonflies A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threate ...
, where males of the Australian emperor dragonfly, ''Hemianax papuensis'' were seen to choose their flight paths to appear stationary to their rivals in 6 of 15 encounters. They made use of both real-point and infinity-point strategies. The strategy appears to work equally well in insects and in vertebrates. Simulations show that motion camouflage results in a more efficient pursuit path than classical pursuit (i.e. the motion camouflage path is shorter), whether the target flies in a straight line or chooses a chaotic path. Further, where classical pursuit requires the attacker to fly faster than the target, the motion camouflaged attacker can sometimes capture the target despite flying more slowly than it. In
sailing Sailing employs the wind—acting on sails, wingsails or kites—to propel a craft on the surface of the ''water'' (sailing ship, sailboat, raft, windsurfer, or kitesurfer), on ''ice'' (iceboat) or on ''land'' (land yacht) over a chosen cour ...
, it has long been known that if the bearing from the target to the pursuer remains constant, known as
constant bearing, decreasing range Constant bearing, decreasing range (CBDR) is a term in navigation which means that some object, usually another ship viewed from the deck or bridge of one's own ship, is getting closer but maintaining the same absolute bearing. If this continues ...
(CBDR), equivalent to taking a fixed reference point at infinite distance, the two vessels are on a collision course, both travelling in straight lines. In a simulation, this is readily observed as the lines between the two remain parallel at all times. Echolocating
bat Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
s follow an infinity-point path when hunting insects in the dark. This is not for camouflage but for the efficiency of the resulting path, so the strategy is generally called constant absolute target direction (CATD); it is equivalent to CBDR but allowing for the target to manoeuvre erratically. A 2014 study of falcons of different species (
gyrfalcon The gyrfalcon ( or ) (), the largest of the falcon species, is a bird of prey. The abbreviation gyr is also used. It breeds on Arctic coasts and tundra, and the islands of northern North America and the Eurosiberian region. It is mainly a reside ...
,
saker falcon The saker falcon (''Falco cherrug'') is a large species of falcon. This species breeds from central Europe eastwards across the Palearctic to Manchuria. It is mainly migratory except in the southernmost parts of its range, wintering in Ethiopia ...
, and
peregrine falcon The peregrine falcon (''Falco peregrinus''), also known as the peregrine, and historically as the duck hawk in North America, is a cosmopolitan bird of prey ( raptor) in the family Falconidae. A large, crow-sized falcon, it has a blue-grey ba ...
) used
video camera A video camera is an optical instrument that captures videos (as opposed to a movie camera, which records images on film). Video cameras were initially developed for the television industry but have since become widely used for a variety of othe ...
s mounted on their heads or backs to track their approaches to prey. Comparison of the observed paths with
simulation A simulation is the imitation of the operation of a real-world process or system over time. Simulations require the use of models; the model represents the key characteristics or behaviors of the selected system or process, whereas the ...
s of different pursuit strategies showed that these predatory birds used a motion camouflage path consistent with CATD. The missile guidance strategy of pure
proportional navigation image:Navigation Constant illustration.png, For example, if the line of sight rotates slowly from north to east, the missile should turn to the right by a certain factor faster than the LOS-rate. This factor is ''N''. Proportional navigation (also ...
guidance (PPNG) closely resembles the CATD strategy used by bats. The biologists Andrew Anderson and Peter McOwan have suggested that anti-aircraft missiles could exploit motion camouflage to reduce their chances of being detected. They tested their ideas on people playing a computerised war game. The steering laws to achieve motion camouflage have been analysed mathematically. The resulting paths turn out to be extremely efficient, often better than classical pursuit. Motion camouflage pursuit may therefore be adopted both by predators and missile engineers (as "parallel navigation", for an infinity-point algorithm) for its performance advantages.


Camouflage by motion


Swaying: motion crypsis or masquerade

Swaying behaviour is practised by highly cryptic animals such as the
leafy sea dragon The leafy seadragon (''Phycodurus eques'') or Glauert's seadragon, is the only member of the genus ''Phycodurus'' and is a marine fish in the family Syngnathidae, which includes seadragons, pipefish, and seahorses. It is found along the sou ...
, the
stick insect The Phasmatodea (also known as Phasmida, Phasmatoptera or Spectra) are an order of insects whose members are variously known as stick insects, stick-bugs, walking sticks, stick animals, or bug sticks. They are also occasionally referred to as ...
''
Extatosoma tiaratum ''Extatosoma tiaratum'', commonly known as the spiny leaf insect, the giant prickly stick insect, Macleay's spectre, or the Australian walking stick, is a large species of Australian stick insect endemic to Australia. The species has the Phasmi ...
'', and
mantis Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They ha ...
es. These animals resemble vegetation with their coloration, strikingly disruptive body outlines with leaflike appendages, and the ability to sway effectively like the plants that they mimic. ''E. tiaratum'' actively sways back and forth or side to side when disturbed or when there is a gust of
wind Wind is the natural movement of air or other gases relative to a planet's surface. Winds occur on a range of scales, from thunderstorm flows lasting tens of minutes, to local breezes generated by heating of land surfaces and lasting a few hou ...
, with a
frequency Frequency is the number of occurrences of a repeating event per unit of time. It is also occasionally referred to as ''temporal frequency'' for clarity, and is distinct from ''angular frequency''. Frequency is measured in hertz The hertz ...
distribution like foliage rustling in the wind. This behaviour may represent motion crypsis, preventing detection by predators, or motion masquerade, promoting misclassification (as something other than prey), or a combination of the two, and has accordingly also been described as a form of motion camouflage.


References


External links

{{Camouflage Biological defense mechanisms Camouflage Antipredator adaptations