Animal locomotion

Source: Wikipedia, the free encyclopedia.
For the Eadweard Muybridge study, see Animal Locomotion.
beetle larva performing a rectilinear locomotion
.

Animal locomotion, in

phoresis
).

Animals move for a variety of reasons, such as to find

microhabitat, or to escape predators. For many animals, the ability to move is essential for survival and, as a result, natural selection has shaped the locomotion methods and mechanisms used by moving organisms. For example, migratory animals that travel vast distances (such as the Arctic tern) typically have a locomotion mechanism that costs very little energy per unit distance, whereas non-migratory animals that must frequently move quickly to escape predators
are likely to have energetically costly, but very fast, locomotion.

The anatomical structures that animals use for movement, including cilia, legs, wings, arms, fins, or tails are sometimes referred to as locomotory organs[2] or locomotory structures.[3]

Etymology

The term "locomotion" is formed in English from Latin loco "from a place" (ablative of locus "place") + motio "motion, a moving".[4]

Locomotion in different media

Animals move through, or on, five types of environment: aquatic (in or on water), terrestrial (on ground or other surface, including arboreal, or tree-dwelling), fossorial (underground), and aerial (in the air). Many animals—for example semi-aquatic animals, and diving birds—regularly move through more than one type of medium. In some cases, the surface they move on facilitates their method of locomotion.

Aquatic

Main article: Aquatic locomotion

Swimming

Dolphins surfing
Further information: Fish locomotion

In water, staying afloat is possible using buoyancy. If an animal's body is less dense than water, it can stay afloat. This requires little energy to maintain a vertical position, but requires more energy for locomotion in the horizontal plane compared to less buoyant animals. The drag encountered in water is much greater than in air. Morphology is therefore important for efficient locomotion, which is in most cases essential for basic functions such as catching prey. A fusiform, torpedo-like body form is seen in many aquatic animals,[5][6] though the mechanisms they use for locomotion are diverse.

The primary means by which

labriform swimming. Marine mammals
oscillate their body in an up-and-down (dorso-ventral) direction. Other animals, e.g. penguins, diving ducks, move underwater in a manner which has been termed "aquatic flying".[7] Some fish propel themselves without a wave motion of the body, as in the slow-moving seahorses and Gymnotus.[8]

Other animals, such as

cetaceans
, now very distinct from their terrestrial ancestors.

Dolphins sometimes ride on the bow waves created by boats or surf on naturally breaking waves.[10]

Benthic

Scallop in jumping motion; these bivalves can also swim.

Echinoderms primarily use their tube feet to move about. The tube feet typically have a tip shaped like a suction pad that can create a vacuum through contraction of muscles. This, along with some stickiness from the secretion of mucus, provides adhesion. Waves of tube feet contractions and relaxations move along the adherent surface and the animal moves slowly along.[11] Some sea urchins also use their spines for benthic locomotion.[12]

Crabs typically walk sideways[13] (a behaviour that gives us the word crabwise). This is because of the articulation of the legs, which makes a sidelong gait more efficient.[14] However, some crabs walk forwards or backwards, including raninids,[15] Libinia emarginata[16] and Mictyris platycheles.[13] Some crabs, notably the Portunidae and Matutidae, are also capable of swimming,[17] the Portunidae especially so as their last pair of walking legs are flattened into swimming paddles.[18]

A stomatopod, Nannosquilla decemspinosa, can escape by rolling itself into a self-propelled wheel and somersault backwards at a speed of 72 rpm. They can travel more than 2 m using this unusual method of locomotion.[19]

Aquatic Surface

Velella moves by sailing.
Main article: Animal locomotion on the water surface

aerofoil, so that the animals tend to sail downwind at a small angle to the wind.[20]

While larger animals such as ducks can move on water by floating, some small animals move across it without breaking through the surface. This surface locomotion takes advantage of the

basilisk lizard.[22]

Aerial

See also: Aeroplankton

Active flight

A pair of brimstone butterflies in flight. The female, above, is in fast forward flight with a small angle of attack; the male, below, is twisting his wings sharply upward to gain lift and fly up towards the female.
Main articles: Flight and Flying and gliding animals

Gravity is the primary obstacle to

better source needed
]

Gliding

Main article: Flying and gliding animals

Rather than active flight, some (semi-) arboreal animals reduce their rate of falling by

gliding. Gliding is heavier-than-air flight without the use of thrust; the term "volplaning" also refers to this mode of flight in animals.[30] This mode of flight involves flying a greater distance horizontally than vertically and therefore can be distinguished from a simple descent like a parachute. Gliding has evolved on more occasions than active flight. There are examples of gliding animals in several major taxonomic classes such as the invertebrates (e.g., gliding ants), reptiles (e.g., banded flying snake), amphibians (e.g., flying frog), mammals (e.g., sugar glider, squirrel glider
).

Flying fish taking off

Some aquatic animals also regularly use gliding, for example, flying fish, octopus and squid. The flights of flying fish are typically around 50 meters (160 ft),[31] though they can use updrafts at the leading edge of waves to cover distances of up to 400 m (1,300 ft).[31][32] To glide upward out of the water, a flying fish moves its tail up to 70 times per second.[33] Several oceanic

Pacific flying squid, leap out of the water to escape predators, an adaptation similar to that of flying fish.[34] Smaller squids fly in shoals, and have been observed to cover distances as long as 50 m. Small fins towards the back of the mantle help stabilize the motion of flight. They exit the water by expelling water out of their funnel, indeed some squid have been observed to continue jetting water while airborne providing thrust even after leaving the water. This may make flying squid the only animals with jet-propelled aerial locomotion.[35] The neon flying squid has been observed to glide for distances over 30 m, at speeds of up to 11.2 m/s.[36]

Soaring

Soaring birds can maintain flight without wing flapping, using rising air currents. Many gliding birds are able to "lock" their extended wings by means of a specialized tendon.

lee waves, convergences and dynamic soaring
.

Examples of soaring flight by birds are the use of:

Ballooning

gossamer silk for ballooning, sometimes traveling great distances at high altitude.[40][41]

Terrestrial

Main article: Terrestrial locomotion
See also: Comparative foot morphology
Pacific leaping blenny Alticus arnoldorum hopping
Pacific leaping blenny climbing up a vertical piece of Plexiglas

Forms of locomotion on land include walking, running, hopping or

bipedal animals, standing on two feet and keeping one on the ground at all times while walking. When running, only one foot is on the ground at any one time at most, and both leave the ground briefly. At higher speeds momentum
helps keep the body upright, so more energy can be used in movement.

Jumping

Main article: Jumping
Gray squirrel (Sciurus carolinensis) in mid-leap

Jumping (saltation) can be distinguished from running, galloping, and other gaits where the entire body is temporarily airborne by the relatively long duration of the aerial phase and high angle of initial launch. Many terrestrial animals use jumping (including hopping or leaping) to escape predators or catch prey—however, relatively few animals use this as a primary mode of locomotion. Those that do include the

Litoria nasuta, can leap over 2 metres (6 ft 7 in), more than fifty times its body length.[47]

Leech moving by looping using its front and back suckers

Peristalsis and looping

Other animals move in terrestrial habitats without the aid of legs. Earthworms crawl by a peristalsis, the same rhythmic contractions that propel food through the digestive tract.[48]

Leech moving on a flat surface

Leeches and geometer moth caterpillars move by looping or inching (measuring off a length with each movement), using their paired circular and longitudinal muscles (as for peristalsis) along with the ability to attach to a surface at both anterior and posterior ends. One end is attached, often the thicker end, and the other end, often thinner, is projected forward peristaltically until it touches down, as far as it can reach; then the first end is released, pulled forward, and reattached; and the cycle repeats. In the case of leeches, attachment is by a sucker at each end of the body.[49]

Sliding

Due to its low coefficient of friction, ice provides the opportunity for other modes of locomotion. Penguins either waddle on their feet or slide on their bellies across the snow, a movement called tobogganing, which conserves energy while moving quickly. Some pinnipeds perform a similar behaviour called sledding.

Climbing

Some animals are specialized for moving on non-horizontal surfaces. One common habitat for such

arboreal movement, travelling rapidly by brachiation (see below
).

Others living on rock faces such as in

rocky mountain goat, etc.), whose adaptations can include a soft rubbery pad between their hooves for grip, hooves with sharp keratin rims for lodging in small footholds, and prominent dew claws. Another case is the snow leopard
, which being a predator of such caprids also has spectacular balance and leaping abilities, such as ability to leap up to 17 m (50 ft).

Some light animals are able to climb up smooth sheer surfaces or hang upside down by adhesion using suckers. Many insects can do this, though much larger animals such as geckos can also perform similar feats.

Walking and running

Species have different numbers of legs resulting in large differences in locomotion.

Modern birds, though classified as

homininan apes. Bipedalism is rarely found outside terrestrial animals—though at least two types of octopus walk bipedally on the sea floor using two of their arms, so they can use the remaining arms to camouflage themselves as a mat of algae or floating coconut.[51]

There are no three-legged animals—though some macropods, such as kangaroos, that alternate between resting their weight on their muscular tails and their two hind legs could be looked at as an example of tripedal locomotion in animals.

Animation of a Devonian tetrapod

Many familiar animals are quadrupedal, walking or running on four legs. A few birds use quadrupedal movement in some circumstances. For example, the shoebill sometimes uses its wings to right itself after lunging at prey.[52] The newly hatched hoatzin bird has claws on its thumb and first finger enabling it to dexterously climb tree branches until its wings are strong enough for sustained flight.[53] These claws are gone by the time the bird reaches adulthood.

A relatively few animals use five limbs for locomotion.

Prehensile
quadrupeds may use their tail to assist in locomotion and when grazing, the kangaroos and other macropods use their tail to propel themselves forward with the four legs used to maintain balance.

Insects generally walk with six legs—though some insects such as nymphalid butterflies[54] do not use the front legs for walking.

harvestmen extend their knees by the use of highly elastic thickenings in the joint cuticle.[55] Scorpions, pseudoscorpions and some harvestmen have evolved muscles that extend two leg joints (the femur-patella and patella-tibia joints) at once.[56][57]

The scorpion Hadrurus arizonensis walks by using two groups of legs (left 1, right 2, Left 3, Right 4 and Right 1, Left 2, Right 3, Left 4) in a reciprocating fashion. This alternating tetrapod coordination is used over all walking speeds.[58]

Centipedes and millipedes have many sets of legs that move in

sunflower seastar (Pycnopodia helianthoides) pull themselves along with some of their arms while letting others trail behind. Other starfish turn up the tips of their arms while moving, which exposes the sensory tube feet and eyespot to external stimuli.[62] Most starfish cannot move quickly, a typical speed being that of the leather star (Dermasterias imbricata), which can manage just 15 cm (6 in) in a minute.[63] Some burrowing species from the genera Astropecten and Luidia have points rather than suckers on their long tube feet and are capable of much more rapid motion, "gliding" across the ocean floor. The sand star (Luidia foliolata) can travel at a speed of 2.8 m (9 ft 2 in) per minute.[64] Sunflower starfish are quick, efficient hunters, moving at a speed of 1 m/min (3.3 ft/min) using 15,000 tube feet.[65]

Many animals temporarily change the number of legs they use for locomotion in different circumstances. For example, many quadrupedal animals switch to bipedalism to reach low-level browse on trees. The genus of

Basiliscus are arboreal lizards that usually use quadrupedalism in the trees. When frightened, they can drop to water below and run across the surface on their hind limbs at about 1.5 m/s for a distance of approximately 4.5 m (15 ft) before they sink to all fours and swim. They can also sustain themselves on all fours while "water-walking" to increase the distance travelled above the surface by about 1.3  m.[66] When cockroaches run rapidly, they rear up on their two hind legs like bipedal humans; this allows them to run at speeds up to 50 body lengths per second, equivalent to a "couple hundred miles per hour, if you scale up to the size of humans."[67]
When grazing, kangaroos use a form of pentapedalism (four legs plus the tail) but switch to hopping (bipedalism) when they wish to move at a greater speed.

  • Bipedal ostrich
  • Hexapedal stick-insect
  • Octopedal locomotion by a spider
  • Multi-legged millipede

Powered cartwheeling

The Moroccan flic-flac spider (

cartwheeling as a form of locomotion.[69] The flic-flac spider can reach speeds of up to 2 m/s using forward or back flips to evade threats.[70][71]

Subterranean

Some animals move through solids such as soil by burrowing using

ground squirrels, naked mole-rats, tilefish, and mole crickets
.

Arboreal locomotion

Main article: arboreal locomotion
A brachiating gibbon

Arboreal locomotion is the locomotion of animals in trees. Some animals may only scale trees occasionally, while others are exclusively arboreal. These habitats pose numerous mechanical challenges to animals moving through them, leading to a variety of anatomical, behavioural and ecological consequences as well as variations throughout different species.

synapsid of the late Permian, about 260 million years ago.[74] Some invertebrate animals are exclusively arboreal in habitat, for example, the tree snail
.

Brachiation (from brachium, Latin for "arm") is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms. During brachiation, the body is alternately supported under each forelimb. This is the primary means of locomotion for the small gibbons and siamangs of southeast Asia. Some New World monkeys such as spider monkeys and muriquis are "semibrachiators" and move through the trees with a combination of leaping and brachiation. Some New World species also practice suspensory behaviors by using their prehensile tail, which acts as a fifth grasping hand.[75]

Pandas are known to swig their heads laterally as they ascend vertical surfaces astonishingly utilizing their head as a propulsive limb in a anatomical way that was thought to only be practiced by certain species of birds.

Energetics

Animal locomotion requires

aerodynamically efficient body shapes of flying birds
indicate how they have evolved to cope with this. Limbless organisms moving on land must energetically overcome surface friction, however, they do not usually need to expend significant energy to counteract gravity.

air).[76] The effect of forces during locomotion on the design of the skeletal system is also important, as is the interaction between locomotion and muscle physiology, in determining how the structures and effectors of locomotion enable or limit animal movement. The energetics of locomotion involves the energy expenditure by animals in moving. Energy consumed in locomotion is not available for other efforts, so animals typically have evolved to use the minimum energy possible during movement.[76]
However, in the case of certain behaviors, such as locomotion to escape a predator, performance (such as speed or maneuverability) is more crucial, and such movements may be energetically expensive. Furthermore, animals may use energetically expensive methods of locomotion when environmental conditions (such as being within a burrow) preclude other modes.

The most common metric of energy use during locomotion is the net (also termed "incremental") cost of transport, defined as the amount of energy (e.g.,

rodents, such as deer mice, the cost of transport has also been measured during voluntary wheel running.[77]

Energetics is important for explaining the evolution of foraging economic decisions in organisms; for example, a study of the African honey bee,

A. m. scutellata, has shown that honey bees may trade the high sucrose content of viscous nectar off for the energetic benefits of warmer, less concentrated nectar, which also reduces their consumption and flight time.[78]

Passive locomotion

Passive locomotion in animals is a type of mobility in which the animal depends on their environment for transportation; such animals are vagile but not motile.[1]

Hydrozoans

Physalia physalis

The Portuguese man o' war (Physalia physalis) lives at the surface of the ocean. The gas-filled bladder, or pneumatophore (sometimes called a "sail"), remains at the surface, while the remainder is submerged. Because the Portuguese man o' war has no means of propulsion, it is moved by a combination of winds, currents, and tides. The sail is equipped with a siphon. In the event of a surface attack, the sail can be deflated, allowing the organism to briefly submerge.[79]

Mollusca

The

mucins to float at the sea surface.[80][81]

Arachnids

The

Namib Desert of Southern Africa. The spider escapes parasitic pompilid wasps by flipping onto its side and cartwheeling down sand dunes at speeds of up to 44 turns per second.[82][83] If the spider is on a sloped dune, its rolling speed may be 1 metre per second.[84]

A spider (usually limited to individuals of a small species), or spiderling after hatching,

spinnerets into the air. These form a triangle-shaped parachute that carries the spider on updrafts of winds, where even the slightest breeze transports it. The Earth's static electric field may also provide lift in windless conditions.[87]

Insects

The

Cicindela dorsalis, the eastern beach tiger beetle, is notable for its ability to leap into the air, loop its body into a rotating wheel and roll along the sand at a high speed using wind to propel itself. If the wind is strong enough, the larva can cover up to 60 metres (200 ft) in this manner. This remarkable ability may have evolved to help the larva escape predators such as the thynnid wasp Methocha.[88]

Members of the largest subfamily of cuckoo wasps,

kleptoparasites
, laying their eggs in host nests, where their larvae consume the host egg or larva while it is still young. Chrysidines are distinguished from the members of other subfamilies in that most have flattened or concave lower abdomens and can curl into a defensive ball when attacked by a potential host, a process known as conglobation. Protected by hard chitin in this position, they are expelled from the nest without injury and can search for a less hostile host.

Fleas can jump vertically up to 18 cm and horizontally up to 33 cm;[89] however, although this form of locomotion is initiated by the flea, it has little control of the jump—they always jump in the same direction, with very little variation in the trajectory between individual jumps.[90][91]

Crustaceans

Although

lobsters, one species, Nannosquilla decemspinosa, has been observed flipping itself into a crude wheel. The species lives in shallow, sandy areas. At low tides, N. decemspinosa is often stranded by its short rear legs, which are sufficient for locomotion when the body is supported by water, but not on dry land. The mantis shrimp then performs a forward flip in an attempt to roll towards the next tide pool. N. decemspinosa has been observed to roll repeatedly for 2 m (6.6 ft), but they typically travel less than 1 m (3.3 ft). Again, the animal initiates the movement but has little control during its locomotion.[92]

Animal transport

Main article: Phoresis

Some animals change location because they are attached to, or reside on, another animal or moving structure. This is arguably more accurately termed "animal transport".

Remoras

Echeneis naucrates
, may attach themselves to scuba divers.

Remoras are a family (Echeneidae) of ray-finned fish.[93][94] They grow to 30–90 cm (0.98–2.95 ft) long, and their distinctive first dorsal fins take the form of a modified oval, sucker-like organ with slat-like structures that open and close to create suction and take a firm hold against the skin of larger marine animals.[95] By sliding backward, the remora can increase the suction, or it can release itself by swimming forward. Remoras sometimes attach to small boats. They swim well on their own, with a sinuous, or curved, motion. When the remora reaches about 3 cm (1.2 in), the disc is fully formed and the remora can then attach to other animals. The remora's lower jaw projects beyond the upper, and the animal lacks a swim bladder. Some remoras associate primarily with specific host species. They are commonly found attached to sharks, manta rays, whales, turtles, and dugongs. Smaller remoras also fasten onto fish such as tuna and swordfish, and some small remoras travel in the mouths or gills of large manta rays, ocean sunfish, swordfish, and sailfish. The remora benefits by using the host as transport and protection, and also feeds on materials dropped by the host.

Angler fish

In some species of anglerfish, when a male finds a female, he bites into her skin, and releases an enzyme that digests the skin of his mouth and her body, fusing the pair down to the blood-vessel level. The male becomes dependent on the female host for survival by receiving nutrients via their shared circulatory system, and provides sperm to the female in return. After fusing, males increase in volume and become much larger relative to free-living males of the species. They live and remain reproductively functional as long as the female lives, and can take part in multiple spawnings. This extreme sexual dimorphism ensures, when the female is ready to spawn, she has a mate immediately available. Multiple males can be incorporated into a single individual female with up to eight males in some species, though some taxa appear to have a one male per female rule.[96][97]

Parasites

Many

phoresis) and attempt to find a new host.[98]

Changes between media

Some animals locomote between different media, e.g., from aquatic to aerial. This often requires different modes of locomotion in the different media and may require a distinct transitional locomotor behaviour.

There are a large number of semi-aquatic animals (animals that spend part of their life cycle in water, or generally have part of their anatomy underwater). These represent the major taxa of mammals (e.g., beaver, otter, polar bear), birds (e.g., penguins, ducks), reptiles (e.g., anaconda, bog turtle, marine iguana) and amphibians (e.g., salamanders, frogs, newts).

Fish

Some fish use multiple modes of locomotion.

mudskippers and the walking catfish. Flying fish can make powerful, self-propelled leaps out of water into air, where their long, wing-like fins enable gliding flight for considerable distances above the water's surface. This uncommon ability is a natural defence mechanism to evade predators. The flights of flying fish are typically around 50 m,[31] though they can use updrafts at the leading edge of waves to cover distances of up to 400 m (1,300 ft).[31][32] They can travel at speeds of more than 70 km/h (43 mph).[33] Maximum altitude is 6 m (20 ft) above the surface of the sea.[99] Some accounts have them landing on ships' decks.[33][100]

Marine mammals

Pacific white-sided dolphins porpoising

When swimming, several marine mammals such as dolphins, porpoises and pinnipeds, frequently leap above the water surface whilst maintaining horizontal locomotion. This is done for various reasons. When travelling, jumping can save dolphins and porpoises energy as there is less friction while in the air.

parasites.[102] In pinnipeds, two types of porpoising have been identified. "High porpoising" is most often near (within 100 m) the shore and is often followed by minor course changes; this may help seals get their bearings on beaching or rafting sites. "Low porpoising" is typically observed relatively far (more than 100 m) from shore and often aborted in favour of anti-predator movements; this may be a way for seals to maximize sub-surface vigilance and thereby reduce their vulnerability to sharks[103]

Some whales raise their (entire) body vertically out of the water in a behaviour known as "breaching".

Birds

Some semi-aquatic birds use terrestrial locomotion, surface swimming, underwater swimming and flying (e.g., ducks, swans).

ratites) have lost the primary locomotion of flight. The largest of these, ostriches, when being pursued by a predator, have been known to reach speeds over 70 km/h (43 mph),[104] and can maintain a steady speed of 50 km/h (31 mph), which makes the ostrich the world's fastest two-legged animal:[105][106] Ostriches can also locomote by swimming.[107]
Penguins either waddle on their feet or slide on their bellies across the snow, a movement called tobogganing, which conserves energy while moving quickly. They also jump with both feet together if they want to move more quickly or cross steep or rocky terrain. To get onto land, penguins sometimes propel themselves upwards at a great speed to leap out the water.

Changes during the life-cycle

An animal's mode of locomotion may change considerably during its life-cycle.

Barnacles are exclusively marine and tend to live in shallow and tidal waters. They have two nektonic (active swimming) larval stages, but as adults, they are sessile
(non-motile) suspension feeders. Frequently, adults are found attached to moving objects such as whales and ships, and are thereby transported (passive locomotion) around the oceans.

Function

Paddlefish ram suspension-feeding zooplankton in aquarium

Animals locomote for a variety of reasons, such as to find food, a mate, a suitable microhabitat, or to escape predators.

Food procurement

Animals use locomotion in a wide variety of ways to procure food. Terrestrial methods include

filterfeeding, grazing, ram feeding, suction feeding, protrusion and pivot feeding. Other methods include parasitism and parasitoidism
.

Quantifying body and limb movement

Main article: Study of animal locomotion

The study of animal locomotion is a branch of biology that investigates and quantifies how animals move. It is an application of kinematics, used to understand how the movements of animal limbs relate to the motion of the whole animal, for instance when walking or flying.[108][109][110]

Galleries

Swimming in major groups of formerly terrestrial animals
Flight in major groups
  • Australian Emperor dragonfly (Insecta)
    Australian Emperor dragonfly (Insecta)
  • Magpie goose (Aves)
    Magpie goose (Aves)
  • Townsend's big-eared bat (Chiroptera)
    Townsend's big-eared bat (Chiroptera)

See also

References

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Further reading

External links

Media related to Animal locomotion at Wikimedia Commons

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