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The prevailing type of fish locomotion is swimming in water. In addition, some fish can
"walk", i.e., move over land, burrow in mud, and glide through the air.

Contents
[hide]

1 Swimming
o 1.1 Body/caudal fin propulsion

1.1.1 Anguilliform locomotion

1.1.2 Sub-carangiform locomotion

1.1.3 Carangiform locomotion

1.1.4 Thunniform locomotion

1.1.5 Ostraciiform locomotion

o 1.2 Median/paired fin propulsion

o 1.3 Dynamic lift
o 1.4 Hydrodynamic principles

1.4.1 Median-paired fin

1.4.1.1 Undulatory

1.4.1.2 Oscillatory

1.4.2 Body-caudal fin

1.4.2.1 Undulatory

1.4.2.2 Oscillatory

o 1.5 Adaptation

2 Flying
o 2.1 Tradeoffs
o 2.2 Biplane body plan
o 2.3 Monoplane body plan

3 Walking

4 Burrowing

5 See also

6 References

7 Further reading

8 External links

Swimming[edit]
Fish swim by exerting force against the surrounding water. There are exceptions, but this is
normally achieved by the fish contracting muscles on either side of its body in order to
generate waves of flexion that travel the length of the body from nose to tail, generally
getting larger as they go along. The vector forces exerted on the water by such motion cancel
out laterally, but generate a net force backwards which in turn pushes the fish forward
through the water.
Most fishes generate thrust using lateral movements of their body and caudal fin. But there
are also a huge number of species that move mainly using their median and paired fins. The
latter group profits from the gained maneuverability that is needed when living in coral reefs
for example. But they can't swim as fast as fish using their bodies and caudal fins.

Body/caudal fin propulsion[edit]

There are five groups that differ in the fraction of their body that is displaced laterally:[1]
Anguilliform locomotion[edit]
In some long, slender fish eels, for example there is little increase in the amplitude of the
flexion wave as it passes along the body.
Sub-carangiform locomotion[edit]
Here, there is a more marked increase in wave amplitude along the body with the vast
majority of the work being done by the rear half of the fish. In general, the fish body is stiffer,
making for higher speed but reduced maneuverability. Trout use sub-carangiform locomotion.

Carangiform locomotion[edit]
Fish in this group, named for the Carangidae, are stiffer and faster-moving than the previous
groups. The vast majority of movement is concentrated in the very rear of the body and tail.
Carangiform swimmers generally have rapidly oscillating tails.
Thunniform locomotion[edit]
The next-to-last group is reserved for the high-speed long-distance swimmers, like tuna (new
research shows that the thunniform locomotion is a unique trait of the tunas[2]). Here, virtually
all the sideways movement is in the tail and the region connecting the main body to the tail
(the peduncle). The tail itself tends to be large and crescent shaped.
Ostraciiform locomotion[edit]
These fishes have no appreciable body wave when they employ caudal locomotion. Only the
tail fin itself oscillates (often very rapidly) to create thrust. This group includes Ostraciidae.

Median/paired fin propulsion[edit]

Not all fish fit comfortably in the above groups. Ocean sunfish, for example, have a
completely different system, and many small fish use their pectoral fins for swimming as well
as for steering and dynamic lift. Fish with electric organs, such as those in Gymnotiformes,
swim by undulating their fins while keeping the body still, presumably so as not to disturb the
electric field that they generate.

Dynamic lift[edit]

Bone and muscle tissues of fish are denser than water. To maintain depth some fish increase
buoyancy by means of a gas bladder or by storing oils or lipids. Fish without these features
use dynamic lift instead. It is done using their pectoral fins in a manner similar to the use of
wings by airplanes and birds. As these fish swim, their pectoral fins are positioned to create
lift which allows the fish to maintain a certain depth.
Sharks are a notable example of fish that depend on dynamic lift; notice their well-developed
pectoral fins.
The two major drawbacks of this method are that these fish must stay moving to stay afloat
and that they are incapable of swimming backwards or hovering.

Hydrodynamic principles[edit]
Similarly to the aerodynamics of flight, powered swimming requires animals to overcome
drag by producing thrust. Unlike flying, however, swimming animals do not necessarily need
to actively exert high vertical forces because the effect of buoyancy can counter the
downward pull of gravity, allowing these animals to float without much effort. While there is
great diversity in fish locomotion, swimming behavior can be classified into two distinct
"modes" based on the body structures involved in thrust production, Median-Paired Fin
(MPF) and Body-Caudal Fin (BCF). Within each of these classifications, there are a
numerous specifications along a spectrum of behaviours from purely undulatory to entirely
oscillatory based. In undulatory swimming modes thrust is produced by wave-like
movements of the propulsive structure (usually a fin or the whole body). Oscillatory modes,
on the other hand, are characterized by thrust production from swiveling of the propulsive
structure on an attachment point without any wave-like motion.[3]

Boxfish are the classic biological example of median-paired fin swimming, because they are
not well streamlined, and use primarily their pectoral fins to produce thrust.
Median-paired fin[edit]
Many fish swim using combined behavior of their two pectoral fins or both their anal and
dorsal fins. Different types of Median paired fin propulsion can be achieved by preferentially
using one fin pair over the other, and include:[3]
Undulatory[edit]

Rajiform: seen in rays, skates, and mantas when thrust is produced by vertical
undulations along large, well developed pectoral fins.

Diodontiform: in which propulsion is achieved by propagating undulations along

large pectoral fins

Amiiform: undulations of a long dorsal fin while the body axis is held straight and
stable

Gymnotiform: undulations of a long anal fin, essentially upside down amiiform

Balistiform: both anal and dorsal fins undulate

Oscillatory[edit]

Tetradontiform:dorsal and anal fins are flapped as a unit, either in phase or exactly
opposing one another. The ocean sunfish is an extreme example of this form of
locomotion.

Labriform: oscillatory movements of pectoral fins and can be classified as drag

based or lift based in which propulsion is generated either as a reaction to drag
produced by dragging the fins through the water in a rowing motion or via lift
mechanisms.[4]
Play media

Sardines use body-caudal fin propulsion to swim and hold their pectoral, dorsal, and anal fins
flat against the body, creating a more streamlined body and reducing drag.
Body-caudal fin[edit]
Most fish swim by generating undulatory waves that propagate down the body through the
caudal fin. This form of undulatory locomotion is termed Body-Caudal Fin (BCF) swimming
on the basis of the body structures used.[3][5]
Undulatory[edit]

Anguilliform: seen in eels and lampreys, this locomotion mode is marked by whole
body in large amplitude wavelengths. Both forward and backward swimming is
possible in this type of BCF swimming.[6]

Subcarangiform: similar to anguilliform swimming, but with limited amplitude

anteriorly that increases as the wave propagates posteriorly, this locomotion mode is
often seen in trout.

Carangiform: body undulations are restricted to the posterior third of body length
with thrust produced by a stiff caudal fin

Thunniform: the most efficient aquatic locomotion mode with thrust being generated
by lift during the lateral movements occurring in the caudal fin only. this locomotion
mode has evolved under independent circumstances in teleost (ray-finned) fish,
sharks, and marine mammals.

Oscillatory[edit]

Ostraciiform: the body remains rigid and the stiff caudal fin is swept in a pendulumlike oscillation. Fish using this type of BCF locomotion, usually rely predominantly
on MPF swimming modes, with ostraciiform behavior only an auxiliary behavior.

Adaptation[edit]
Similar to adaptation in avian flight, swimming behaviors in fish can be thought of as a
balance of stability and maneuverability.[7] Because BCF swimming relies on more caudal
body structures that can direct powerful thrust only rearwards, this form of locomotion is

particularly effective for accelerating quickly and cruising continuously.[3][5] BCF swimming
is, therefore, inherently stable and is often seen in fish with large migration patterns that must
maximize efficiency over long periods. Propulsive forces in MPF swimming, on the other
hand, are generated by multiple fins located on either side of the body that can be coordinated
to execute elaborate turns. As a result, MPF swimming is well adapted for high
maneuverability and is often seen in smaller fish that require elaborate escape patterns.[7]
The habitats occupied by fishes are often related to their swimming capabilities. Coral reefs
typically have faster-swimming fish species living in wave-swept habitats subject to fast
water flow speeds, while the slower fishes live in sheltered habitats with low levels of water
movement.[8]
It is important to point out that fish do not rely exclusively on one locomotor mode, but are
rather locomotor "generalists,"[3] choosing among and combining behaviors from many
available behavioral techniques. In fact, at slower speeds, predominantly BCF swimmers will
often incorporate movement of their pectoral, anal, and dorsal fins as an additional stabilizing
mechanism at slower speeds,[9] but hold them close to their body at high speeds to improve
streamlining and reducing drag.[3] Zebrafish have even been observed to alter their locomotor
behavior in response to changing hydrodynamic influences throughout growth and
maturation.[10]
In addition to adapting locomotor behavior, controlling buoyancy effects is critical for aquatic
survival since aquatic ecosystems vary greatly by depth. Fish generally control their depth by
regulating the amount of gas in specialized organs that are much like balloons. By changing
the amount of gas in these swim bladders, fish actively control their density. If they increase
the amount of air in their swim bladder, their overall density will become less than the
surrounding water, and increased upward buoyancy pressures will cause the fish to rise until
they reach a depth at which they are again at equilibrium with the surrounding water. In this
way, fish behave essentially as a hot air balloon does in air.

Flying[edit]
See also: flying fish and flying and gliding animals
The transition of predominantly swimming locomotion directly to flight has evolved in a
single family of marine fish called Exocoetidae. Flying fish are not true fliers in the sense that
they do not execute powered flight. Instead, these species glide directly over the surface of
the ocean water without ever flapping their "wings." Flying fish have evolved abnormally
large pectoral fins that act as airfoils and provide lift when the fish launches itself out of the
water. Additional forward thrust and steering forces are created by dipping the hypocaudal
(i.e. bottom) lobe of their caudal fin into the water and vibrating it very quickly, in contrast to
diving birds in which these forces are produced by the same locomotor module used for
propulsion. Of the 64 extant species of flying fish, only two distinct body plans exist, each of
which optimizes two different behaviors.[11][12]

Flying fish are able to achieve sufficient lift to glide above the surface of the water thanks to
their enlarged pectoral fins.

Tradeoffs[edit]
Tail Structure: While most fish have caudal fins with evenly sized lobes (i.e. homocaudal),
flying fish have an enlarged ventral lobe (i.e. hypocaudal) which facilitates dipping only a
portion of the tail back onto the water for additional thrust production and steering.[12]
Larger mass: Because flying fish are primarily aquatic animals, their body density must be
close to that of water for buoyancy stability. This primary requirement for swimming,
however, means that flying fish are heavier than other habitual fliers, resulting in higher wing
loading and lift to drag ratios for flying fish compared to a comparably sized bird.[11]
Differences in wing area, wing span, wing loading, and aspect ratio have been used to
classify flying fish into two distinct classifications based on these different aerodynamic
designs.[11]

Biplane body plan[edit]

In the biplane or cypselurus body plan, both the pectoral and pelvic fins are enlarged to
provide lift during flight.[11] These fish also tend to have "flatter" bodies which increase the
total lift producing area thus allowing them to "hang" in the air better than more streamlined
shapes.[12] As a result of this high lift production, these fish are excellent gliders and are well
adapted for maximizing flight distance and duration.
Comparatively, Cypselurus flying fish have lower wing loading and smaller aspect ratios (i.e.
broader wings) than their Exocoetus monoplane counterparts, which contributes to their
ability to fly for longer distances than fish with this alternative body plan. Flying fish with the
biplane design take advantage of their high lift production abilities when launching from the
water by utilizing a "taxiing glide" in which the hypocaudal lobe remains in the water to
generate thrust even after the trunk clears the water's surface and the wings are opened with a
small angle of attack for lift generation.[11]

In the monoplane body plan, only the pectoral fins are abnormally large. In this illustration,
note that the pelvic fins are not abnormally large.

Monoplane body plan[edit]

In the Exocoetus or monoplane body plan, only the pectoral fins are enlarged to provide lift.
Fish with this body plan tend to have a more streamlined body, higher aspect ratios (long,
narrow wings), and higher wing loading than fish with the biplane body plan, making these
fish well adapted for higher flying speeds. Flying fish with a monoplane body plan
demonstrate different launching behaviors from their biplane counterparts. Instead of
extending their duration of thrust production, monoplane fish launch from the water at high
speeds at a large angle of attack (sometimes up to 45 degrees).[11] In this way, monoplane fish
are taking advantage of their adaptation for high flight speed, while fish with biplane designs
exploit their lift production abilities during takeoff.

Walking[edit]
Main article: Walking fish
Play media
Alticus arnoldorum hopping
Play media
Alticus arnoldorum climbing up a vertical piece of Plexiglas
A "walking fish" is a fish that is able to travel over land for extended periods of time. Some
other cases of nonstandard fish locomotion include fish "walking" along the sea floor, such as
the handfish or frogfish.
Most commonly, walking fish are amphibious fish. Able to spend longer times out of water,
these fish may use a number of means of locomotion, including springing, snake-like lateral
undulation, and tripod-like walking. The mudskippers are probably the best land-adapted of
contemporary fish and are able to spend days moving about out of water and can even climb
mangroves, although to only modest heights.[13] The Climbing gourami is often specifically
referred to as a "walking fish", although it does not actually "walk", but rather moves in a
jerky way by supporting itself on the extended edges of its gill plates and pushing itself by its
fins and tail. Some reports indicate that it can also climb trees.[14]
There are a number of fish that are less adept at actual walking, such as the walking catfish.
Despite being known for "walking on land", this fish usually wriggles and may use its
pectoral fins to aid in its movement. Walking Catfish have a respiratory system that allows
them to live out of water for several days. Some are invasive species. A notorious case in the
United States is the Northern snakehead.[15] Polypterids have rudimentary lungs and can also
move about on land, though rather clumsily. The Mangrove rivulus can survive for months
out of water and can move to places like hollow logs.[16][17][18][19]

Ogcocephalus parvus
There are some species of fish that can "walk" along the sea floor but not on land; one such
animal is the flying gurnard (it does not actually fly, and should not be confused with flying
fish). The batfishes of the Ogcocephalidae family (not to be confused with Batfish of
Ephippidae) are also capable of walking along the sea floor. Bathypterois grallator, also
known as a "tripodfish", stands on its three fins on the bottom of the ocean and hunts for
food.[20] The African lungfish (P. annectens) can use its fins to "walk" along the bottom of its
tank in a manner similar to the way amphibians and land vertebrates use their limbs on land.
[21][22][23]

Burrowing[edit]
Many fishes, particularly eel-shaped fishes such as true eels, moray eels, and spiny eels, are
capable of burrowing through sand or mud.[24] Ophichthids are capable of digging backwards
using a sharpened tail.

See also[edit]

Aquatic locomotion

Role of skin in locomotion

Tradeoffs for locomotion in air and water

Undulatory locomotion

References[edit]
1.

Jump up ^ Breder CM (1926) "The locomotion of fishes", Zoologica, 4: 159

297.

2.

Jump up ^ Hawkins JD, CA Sepulveda, JB Graham and KA Dickson (2003)

"Swimming performance studies on the eastern Pacific bonito Sarda chiliensis, a
close relative of the tunas (family Scombridae) II. Kinematics" The Journal of
Experimental Biology, 206: 2749-2758.

3.

^ Jump up to: a b c d e f Sfakiotakis M, Lane DM and Davies JBC (1999)

"Review of Fish Swimming Modes for Aquatic Locomotion" IEEE Journal of
Oceanic Engineering, 24 (2).

4.

Jump up ^ Fulton CJ, Johansen JL, Steffensen JF (2013) Energetic extremes

in aquatic locomotion by coral reef fishes. PLoS One 8, e54033

5.

^ Jump up to: a b Blake, R.W. (2004) Review Paper: Fish functional design and
swimming performance. Journal of Fish Biology 65, pp 1193-1222.

6.

Jump up ^ Long Jr, J. H., Shepherd, W., & Root, R. G. (1997).

Manueuverability and reversible propulsion: How eel-like fish swim forward and
backward using travelling body waves". In: Proc. Special Session on Bio-Engineering
Research Related to Autonomous Underwater Vehicles, 10th Int. Symp. Unmanned
Untethered Submersible Technology (pp. 118-134).

7.

^ Jump up to: a b Weihs, Daniel. (2002) Stability versus Maneuverability in

Aquatic Locomotion. Integrated and Computational Biology. 42, 127-134.

8.

Jump up ^ Fulton CJ, Bellwood DR, Wainwright PC (2005) Wave energy and
swimming performance shape coral reef fish assemblages. Proceedings of the Royal
Society B 272, 827-832

9.

Jump up ^ Heatwole SJ, Fulton CJ (2013) Behavioural flexibility in coral

reef fishes responding to a rapidly changing environment. Marine Biology 160, 677689

10.

Jump up ^ , Matthew J. and George V. Lauder. (2006) Otogeny of Form and

Function: Locomotor Morphology and Drag in Zebrafish (Danio rerio). "Journal of
Morphology." 267,1099-1109.

11.

^ Jump up to: a b c d e f Fish, F.E. (1990) Wing design and scaling of flying fish
with regard to flight performance. "J. Zool. Lond." 221, 391-403.

12.

^ Jump up to: a b c Fish, Frank. (1991) On a Fin and a Prayer. "Scholars." 3(1),
4-7.

13.

Jump up ^ http://www.cairnsmuseum.org.au/tourism.htm

14.

Jump up ^ Climbing Fish

15.
16.
17.

Jump up ^ "Maryland Suffers Setback in War on Invasive Walking Fish",

National Geographic News July 12, 2002
Jump up ^ Shells, trees and bottoms: Strange places fish live
Jump up ^ "Tropical fish can live for months out of water". Reuters. 15
November 2007.

18.

Jump up ^ Fish Lives in Logs, Breathing Air, for Months at a Time

19.

Jump up ^ Fish Lives in Logs, Breathing Air, for Months at a Time

20.

21.

Jump up ^ Jones, AT; KJ Sulak (1990). "First Central Pacific Plate and
Hawaiian Record of the Deep-sea Tripod Fish Bathypterois grallator (Pisces:
Chlorophthalmidae)" (PDF). Pacific Science. 44 (3): 2547.
Jump up ^ Fish uses fins to walk and bound

22.

Jump up ^ Behavioral evidence for the evolution of walking and bounding

before terrestriality in sarcopterygian fishes

23.

Jump up ^ A Small Step for Lungfish, a Big Step for the Evolution of
Walking

24.

Jump up ^ Monks, Neale (2006). Brackish-Water Fishes. TFH. pp. 223226.

ISBN 0-7938-0564-3.

Further reading[edit]

Alexander, R. McNeill (2003) Principles of Animal Locomotion. Princeton University

Press. ISBN 0-691-08678-8.

Eloy, Christophe (2013) "On the best design for undulatory swimming" Journal of
Fluid Mechanics, 717: 4889. doi:10.1017/jfm.2012.561

Lauder GV, Nauen JC and Drucker EG (2002) "Experimental Hydrodynamics and

Evolution: Function of Median Fins in Ray-finned Fishes" Integr. Comp. Biol. 42 (5):
10091017. doi:10.1093/icb/42.5.1009

Videler JJ (1993) Fish Swimming Springer. ISBN 9780412408601.

Vogel, Steven (1994) Life in Moving Fluid: The Physical Biology of Flow. Princeton
University Press. ISBN 0-691-02616-5 (particularly pp. 115117 and pp. 207216 for
specific biological examples swimming and flying respectively)

Wu, Theodore, Y.-T., Brokaw, Charles J., Brennen, Christopher, Eds. (1975)
Swimming and Flying in Nature. Volume 2, Plenum Press. ISBN 0-306-37089-1
(particularly pp. 615652 for an in depth look at fish swimming)

External links[edit]

How fish swim: study solves muscle mystery

Simulated fish locomotion

Basic introduction to the basic principles of biologically inspired swimming robots

The biomechanics of swimming

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