Aquatic Locomotion
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Aquatic locomotion or swimming is biologically propelled motion through a
liquid A liquid is a nearly incompressible fluid that conforms to the shape of its container but retains a (nearly) constant volume independent of pressure. As such, it is one of the four fundamental states of matter (the others being solid, gas, a ...
medium. The simplest propulsive systems are composed of cilia and
flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
. Swimming has evolved a number of times in a range of organisms including
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s,
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
, molluscs,
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s,
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s,
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s, and
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s.


Evolution of swimming

Swimming Swimming is the self-propulsion of a person through water, or other liquid, usually for recreation, sport, exercise, or survival. Locomotion is achieved through coordinated movement of the limbs and the body to achieve hydrodynamic thrust that r ...
evolved a number of times in unrelated lineages. Supposed jellyfish fossils occur in the Ediacaran, but the first free-swimming animals appear in the Early to Middle
Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
. These are mostly related to the
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s, and include the Anomalocaridids, which swam by means of lateral lobes in a fashion reminiscent of today's
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 of ...
. Cephalopods joined the ranks of the
nekton Nekton or necton (from the ) refers to the actively swimming aquatic organisms in a body of water. The term was proposed by German biologist Ernst Haeckel to differentiate between the active swimmers in a body of water, and the passive organisms t ...
in the late Cambrian, and chordates were probably swimming from the Early Cambrian. Many terrestrial animals retain some capacity to swim, however some have returned to the water and developed the capacities for aquatic locomotion. Most apes (including humans), however, lost the swimming
instinct Instinct is the inherent inclination of a living organism towards a particular complex behaviour, containing both innate (inborn) and learned elements. The simplest example of an instinctive behaviour is a fixed action pattern (FAP), in which a v ...
. In 2013 Pedro Renato Bender, a research fellow at the
University of the Witwatersrand The University of the Witwatersrand, Johannesburg (), is a multi-campus South African Public university, public research university situated in the northern areas of central Johannesburg. It is more commonly known as Wits University or Wits ( o ...
's Institute for Human Evolution, proposed a theory to explain the loss of that instinct. Termed the Saci last common ancestor hypothesis (after
Saci ''Sac''I is a restriction enzyme isolated from the bacterium Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorgani ...
, a Brazilian folklore character who cannot cross water barriers), it holds that the loss of instinctive swimming ability in apes is best explained as a consequence of constraints related to the adaptation to an arboreal life in the
last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of apes. Bender hypothesized that the ancestral ape increasingly avoided deep-water bodies when the risks of being exposed to water were clearly higher than the advantages of crossing them. A decreasing contact with water bodies then could have led to the disappearance of the
doggy paddle The dog paddle or doggy paddle is a simple swimming style. It is characterized by the swimmer lying on their chest and moving their hands and legs alternately in a manner reminiscent of how dogs and other quadrupedal mammals swim. It is effectiv ...
instinct.


Micro-organisms

Microbial swimmers, sometimes called
microswimmer A microswimmer is a microscopic object with the ability to move in a fluid environment. Natural microswimmers are found everywhere in the natural world as biological microorganisms, such as bacteria, archaea, protists, sperm and microanimals. Sinc ...
s, are microscopic entities that have the ability to move in fluid or aquatic environment.
Natural microswimmers Nature, in the broadest sense, is the physical world or universe. "Nature" can refer to the phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. Although humans are ...
are found everywhere in the natural world as biological
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s, such as
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
,
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
, protists,
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...
and microanimals.


Bacterial


Ciliates

Ciliates The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a different ...
use small flagella called cilia to move through the water. One ciliate will generally have hundreds to thousands of cilia that are densely packed together in arrays. During movement, an individual cilium deforms using a high-friction power stroke followed by a low-friction recovery stroke. Since there are multiple cilia packed together on an individual organism, they display collective behavior in a
metachronal rhythm A metachronal rhythm or metachronal wave refers to wavy movements produced by the sequential action (as opposed to synchronized) of structures such as cilia, segments of worms, or legs. These movements produce the appearance of a travelling wave. ...
. This means the deformation of one cilium is in phase with the deformation of its neighbor, causing deformation waves that propagate along the surface of the organism. These propagating waves of cilia are what allow the organism to use the cilia in a coordinated manner to move. A typical example of a ciliated microorganism is the ''
Paramecium '' ''Paramecium'' ( , ; also spelled ''Paramoecium'') is a genus of eukaryotic, unicellular ciliates, commonly studied as a representative of the ciliate group. ''Paramecia'' are widespread in freshwater, brackish, and marine environments and a ...
'', a one-celled, ciliated protozoan covered by thousands of cilia. The cilia beating together allow the ''Paramecium'' to propel through the water at speeds of 500 micrometers per second.


Flagellates

Certain organisms such as
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
and animal sperm have
flagellum A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
which have developed a way to move in liquid environments. A rotary motor model shows that bacteria uses the protons of an
electrochemical gradient An electrochemical gradient is a gradient of electrochemical potential, usually for an ion that can move across a membrane. The gradient consists of two parts, the chemical gradient, or difference in solute concentration across a membrane, and ...
in order to move their flagella.
Torque In physics and mechanics, torque is the rotational equivalent of linear force. It is also referred to as the moment of force (also abbreviated to moment). It represents the capability of a force to produce change in the rotational motion of th ...
in the flagella of bacteria is created by particles that conduct protons around the base of the flagellum. The direction of rotation of the flagella in bacteria comes from the occupancy of the proton channels along the perimeter of the flagellar motor. Movement of sperm is called
sperm motility Sperm motility describes the ability of sperm to move properly through the female reproductive tract (internal fertilization) or through water (external fertilization) to reach the egg. Sperm motility can also be thought of as the ''quality'', wh ...
. The middle of the mammalian spermatozoon contains mitochondria that power the movement of the flagellum of the sperm. The motor around the base produces torque, just like in bacteria for movement through the aqueous environment.


Pseudopodia

Movement using a pseudopod is accomplished through increases in pressure at one point on the
cell membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment ( ...
. This pressure increase is the result of
actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of over ...
polymerization between the cortex and the membrane. As the pressure increases the
cell membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment ( ...
is pushed outward creating the pseudopod. When the pseudopod moves outward, the rest of the body is pulled forward by cortical tension. The result is cell movement through the
fluid In physics, a fluid is a liquid, gas, or other material that continuously deforms (''flows'') under an applied shear stress, or external force. They have zero shear modulus, or, in simpler terms, are substances which cannot resist any shear ...
medium. Furthermore, the direction of movement is determined by chemotaxis. When chemoattraction occurs in a particular area of the
cell membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment ( ...
,
actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of over ...
polymerization can begin and move the cell in that direction. An excellent example of an organism that utilizes pseudopods is ''
Naegleria fowleri ''Naegleria fowleri'', colloquially known as a "brain-eating amoeba", is a species of the genus ''Naegleria'', belonging to the phylum Percolozoa, which is technically not classified as true amoeba, but a shapeshifting amoeboflagellate excava ...
.''
A Simple Animation


Invertebrates

Among the
radiata Radiata or Radiates is a historical taxonomic rank that was used to classify animals with radially symmetric body plans. The term Radiata is no longer accepted, as it united several different groupings of animals that do not form a monophyletic ...
, jellyfish and their kin, the main form of swimming is to flex their cup shaped bodies. All
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrella- ...
are free-swimming, although many of these spend most of their time swimming passively. Passive swimming is akin to gliding; the organism floats, using currents where it can, and does not exert any energy into controlling its position or motion. Active swimming, in contrast, involves the expenditure of energy to travel to a desired location. In bilateria, there are many methods of swimming. The
arrow worm The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, about 20% of the known Chaetognatha species are benthic zone, be ...
s (
chaetognatha The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, about 20% of the known Chaetognatha species are benthic, and ca ...
) undulate their finned bodies, not unlike fish.
Nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s swim by undulating their fin-less bodies. Some
Arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
groups can swim - including many
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s. Most crustaceans, such as
shrimp Shrimp are crustaceans (a form of shellfish) with elongated bodies and a primarily swimming mode of locomotion – most commonly Caridea and Dendrobranchiata of the decapod order, although some crustaceans outside of this order are refer ...
, will usually swim by paddling with special swimming legs (
pleopods The decapod ( crustaceans such as a crab, lobster, shrimp or prawn) is made up of 20 body segments grouped into two main body parts: the cephalothorax and the pleon (abdomen). Each segment may possess one pair of appendages, although in variou ...
).
Swimming crab Portunidae is a family of crabs which contains the swimming crabs. Description Portunid crabs are characterised by the flattening of the fifth pair of legs into broad paddles, which are used for swimming. This ability, together with their strong ...
s swim with modified walking legs ( pereiopods).
Daphnia ''Daphnia'' is a genus of small planktonic crustaceans, in length. ''Daphnia'' are members of the order Anomopoda, and are one of the several small aquatic crustaceans commonly called water fleas because their saltatory swimming style resembl ...
, a crustacean, swims by beating its antennae instead. There are also a number of forms of swimming molluscs. Many free-swimming
sea slug Sea slug is a common name for some marine invertebrates with varying levels of resemblance to terrestrial slugs. Most creatures known as sea slugs are gastropods, i.e. they are sea snails (marine gastropod mollusks) that over evolutionary t ...
s, such as
sea angel Sea angels ( clade Gymnosomata) are a large group of small free-swimming sea slugs, not to be confused with Cnidarians (Jellyfish and other similar creatures), classified into six different families. They are pelagic opisthobranchs in the cla ...
s, flap fin-like structures. Some shelled molluscs, such as scallops can briefly swim by clapping their two shells open and closed. The molluscs most evolved for swimming are the
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s. Violet sea-snails exploit a buoyant foam raft stabilized by
amphiphilic An amphiphile (from the Greek αμφις amphis, both, and φιλíα philia, love, friendship), or amphipath, is a chemical compound possessing both hydrophilic (''water-loving'', polar) and lipophilic (''fat-loving'') properties. Such a compo ...
mucins Mucins () are a family of high molecular weight, heavily glycosylated proteins (glycoconjugates) produced by epithelial tissues in most animals. Mucins' key characteristic is their ability to form gels; therefore they are a key component in most ...
to float at the sea surface. Among the
Deuterostomia Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some ex ...
, there are a number of swimmers as well.
Feather star Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which ar ...
s can swim by undulating their many arms.
Salp A salp (plural salps, also known colloquially as “sea grape”) or salpa (plural salpae or salpas) is a barrel-shaped, planktic tunicate. It moves by contracting, thereby pumping water through its gelatinous body, one of the most efficient ...
s move by pumping waters through their gelatinous bodies. The deuterostomes most evolved for swimming are found among the
vertebrate 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, ...
s, notably the
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
.


Jet propulsion

Jet propulsion Jet propulsion is the propulsion of an object in one direction, produced by ejecting a jet of fluid in the opposite direction. By Newton's third law, the moving body is propelled in the opposite direction to the jet. Reaction engines operating o ...
is a method of aquatic locomotion where animals fill a muscular cavity and squirt out water to propel them in the opposite direction of the squirting water. Most organisms are equipped with one of two designs for jet propulsion; they can draw water from the rear and expel it from the rear, such as jellyfish, or draw water from front and expel it from the rear, such as salps. Filling up the cavity causes an increase in both the mass and drag of the animal. Because of the expanse of the contracting cavity, the animal's velocity fluctuates as it moves through the water, accelerating while expelling water and decelerating while vacuuming water. Even though these fluctuations in drag and mass can be ignored if the frequency of the jet-propulsion cycles is high enough, jet-propulsion is a relatively inefficient method of aquatic locomotion. All
cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s can move by
jet propulsion Jet propulsion is the propulsion of an object in one direction, produced by ejecting a jet of fluid in the opposite direction. By Newton's third law, the moving body is propelled in the opposite direction to the jet. Reaction engines operating o ...
, but this is a very energy-consuming way to travel compared to the tail propulsion used by fish. The relative efficiency of jet propulsion decreases further as animal size increases. Since the Paleozoic, as competition with fish produced an environment where efficient motion was crucial to survival, jet propulsion has taken a back role, with fins and tentacles used to maintain a steady velocity. The stop-start motion provided by the jets, however, continues to be useful for providing bursts of high speed - not least when capturing prey or avoiding predators. Indeed, it makes cephalopods the fastest marine invertebrates, and they can out accelerate most fish. Oxygenated water is taken into the
mantle cavity The mantle (also known by the Latin word pallium meaning mantle, robe or cloak, adjective pallial) is a significant part of the anatomy of molluscs: it is the dorsal body wall which covers the visceral mass and usually protrudes in the form of ...
to the
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
s and through muscular contraction of this cavity, the spent water is expelled through the
hyponome A siphon is an anatomical structure which is part of the body of aquatic molluscs in three Class (biology), classes: Gastropoda, Bivalvia and Cephalopoda (members of these classes include saltwater and freshwater snails, clams, octopus, squid and ...
, created by a fold in the mantle. Motion of the cephalopods is usually backward as water is forced out anteriorly through the hyponome, but direction can be controlled somewhat by pointing it in different directions. Most cephalopods float (i.e. are neutrally buoyant), so do not need to swim to remain afloat.
Squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...
swim more slowly than fish, but use more power to generate their speed. The loss in efficiency is due to the amount of water the squid can accelerate out of its mantle cavity.Levinton, Jeffrey S. Marine Biology: Function, Biodiversity, Ecology. 3rd ed. New York, New York: Oxford University Press, 2008. Print.
Jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrella- ...
use a one-way water cavity design which generates a phase of continuous cycles of jet-propulsion followed by a rest phase. The Froude efficiency is about 0.09, which indicates a very costly method of locomotion. The metabolic cost of transport for jellyfish is high when compared to a fish of equal mass. Other jet-propelled animals have similar problems in efficiency. Scallops, which use a similar design to jellyfish, swim by quickly opening and closing their shells, which draws in water and expels it from all sides. This locomotion is used as a means to escape predators such as starfish. Afterwards, the shell acts as a hydrofoil to counteract the scallop's tendency to sink. The Froude efficiency is low for this type of movement, about 0.3, which is why it's used as an emergency escape mechanism from predators. However, the amount of work the scallop has to do is mitigated by the elastic hinge that connects the two shells of the bivalve. Squids swim by drawing water into their mantle cavity and expelling it through their siphon. The Froude efficiency of their jet-propulsion system is around 0.29, which is much lower than a fish of the same mass. Much of the work done by scallop muscles to close its shell is stored as elastic energy in abductin tissue, which acts as a spring to open the shell. The elasticity causes the work done against the water to be low because of the large openings the water has to enter and the small openings the water has to leave. The inertial work of scallop jet-propulsion is also low. Because of the low inertial work, the energy savings created by the elastic tissue is so small that it's negligible. Medusae can also use their elastic mesoglea to enlarge their bell. Their mantle contains a layer of muscle sandwiched between elastic fibers. The muscle fibers run around the bell circumferentially while the elastic fibers run through the muscle and along the sides of the bell to prevent lengthening. After making a single contraction, the bell vibrates passively at the resonant frequency to refill the bell. However, in contrast with scallops, the inertial work is similar to the hydrodynamic work due to how medusas expel water - through a large opening at low velocity. Because of this, the negative pressure created by the vibrating cavity is lower than the positive pressure of the jet, meaning that inertial work of the mantle is small. Thus, jet-propulsion is shown as an inefficient swimming technique.


Fish

Many
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
swim through water by creating undulations with their bodies or oscillating their
fins A fin is a thin component or appendage attached to a larger body or structure. Fins typically function as foils that produce lift or thrust, or provide the ability to steer or stabilize motion while traveling in water, air, or other fluids. Fin ...
. The undulations create components of forward thrust complemented by a rearward force, side forces which are wasted portions of energy, and a normal force that is between the forward thrust and side force. Different fish swim by undulating different parts of their bodies. Eel-shaped fish undulate their entire body in rhythmic sequences. Streamlined fish, such as salmon, undulate the caudal portions of their bodies. Some fish, such as sharks, use stiff, strong fins to create dynamic lift and propel themselves. It is common for fish to use more than one form of propulsion, although they will display one dominant mode of swimming McNeill Alexander, R. Exploring Biomechanics: Animals in Motion. 2nd ed. New York, New York: W. H. Freeman & Co, 1992. Print. Gait changes have even been observed in juvenile reef fish of various sizes. Depending on their needs, fish can rapidly alternate between synchronized fin beats and alternating fin beats.Hale, Melina, R. Day, D. Thorsen, M. Westneat. 2006. Pectoral fin coordination and gait transitions in steadily swimming juvenile reef fishes. The Journal of Experimental Biology (209): 3708-3718. According to ''
Guinness World Records ''Guinness World Records'', known from its inception in 1955 until 1999 as ''The Guinness Book of Records'' and in previous United States editions as ''The Guinness Book of World Records'', is a reference book published annually, listing world ...
2009'', ''
Hippocampus zosterae The dwarf seahorse (''Hippocampus zosterae'') is a species of seahorse found in the subtidal aquatic beds of the Bahamas and parts of the United States. It is threatened by habitat loss. According to ''Guinness World Records'', it is the slowes ...
'' (the dwarf seahorse) is the slowest moving fish, with a top speed of about per hour. They swim very poorly, rapidly fluttering a
dorsal fin A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through conv ...
and using
pectoral fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as ...
s (located behind their eyes) to steer. Seahorses have no
caudal fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
.


Body-caudal fin (BCF) propulsion

*Anguilliform: Anguilliform swimmers are typically slow swimmers. They undulate the majority of their body and use their head as the fulcrum for the load they are moving. At any point during their undulation, their body has an amplitude between 0.5-1.0 wavelengths. The amplitude that they move their body through allows them to swim backwards. Anguilliform locomotion is usually seen in fish with long, slender bodies like
eels Eels are ray-finned fish belonging to the order Anguilliformes (), which consists of eight suborders, 19 families, 111 genera, and about 800 species. Eels undergo considerable development from the early larval stage to the eventual adult stage ...
,
lampreys Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like su ...
,
oarfish Oarfish are huge, greatly elongated, pelagic lampriform fish belonging to the small family Regalecidae. Found in areas spanning from temperate ocean zones to tropical ones, yet rarely seen, the oarfish family contains three species in two gene ...
, and a number of
catfish Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive ...
species. *Subcarangiform, Carangiform, Thunniform: These swimmers undulate the posterior half of their body and are much faster than anguilliform swimmers. At any point while they are swimming, a wavelength <1 can be seen in the undulation pattern of the body. Some Carangiform swimmers include nurse sharks, bamboo sharks, and reef sharks. Thunniform swimmers are very fast and some common Thunniform swimmers include tuna, white sharks, salmon, jacks, and mako sharks. Thunniform swimmers only undulate their high aspect ratio caudal fin, so they are usually very stiff to push more water out of the way. *Ostraciiform: Ostraciiform swimmers oscillate their caudal region, making them relatively slow swimmers. Boxfish, torpedo rays, and momyrs employ Ostraciiform locomotion. The cow fish uses Osctraciiform locomotion to hover in the water column.Sfakiotakis, Michael, D. Lane, B. Davies. 1999. Review of Fish Swimming Modes for Aquatic Locomotion. Journal of Oceanic Engineering (24:2) 237-252.


Median paired fin (MPF) propulsion

*Tetraodoniform, Balistiform, Diodontiform: These swimmers oscillate their median fins. They are typically slow swimmers, and some notable examples include the
oceanic sunfish The ocean sunfish or common mola (''Mola mola'') is one of the largest bony fish in the world. It was misidentified as the heaviest bony fish, which was actually a different species, ''Mola alexandrini''. Adults typically weigh between . The spe ...
(which has extremely modified anal and dorsal fins), puffer fish, and triggerfish. *Rajiform, Amiiform, Gymnotiform: This locomotory mode is accomplished by undulation of the pectoral and median fins. During their undulation pattern, a wavelength >1 can be seen in their fins. They are typically slow to moderate swimmers, and some examples include rays, bowfin, and knife fishes. The
black ghost knife fish The black ghost knifefish (''Apteronotus albifrons'') is a tropical fish belonging to the ghost knifefish family ( Apteronotidae). They originate in freshwater habitats in South America where they range from Venezuela to the Paraguay– Paraná R ...
is a Gymnotiform swimmer that has a very long ventral ribbon fin. Thrust is produced by passing waves down the ribbon fin while the body remains rigid. This also allows the ghost knife fish to swim in reverse. *Labriform: Labriform swimmers are also slow swimmers. They oscillate their pectoral fins to create thrust. Oscillating fins create thrust when a starting vortex is shed from the trailing edge of the fin. As the foil departs from the starting vortex, the effect of that vortex diminishes, while the bound circulation remains, producing lift. Labriform swimming can be viewed as continuously starting and stopping.
Wrasses The wrasses are a family, Labridae, of marine fish, many of which are brightly colored. The family is large and diverse, with over 600 species in 81 genera, which are divided into 9 subgroups or tribes. They are typically small, most of them le ...
and surf perch are common Labriform swimmers.


Hydrofoils

Hydrofoils A hydrofoil is a lifting surface, or foil, that operates in water. They are similar in appearance and purpose to aerofoils used by aeroplanes. Boats that use hydrofoil technology are also simply termed hydrofoils. As a hydrofoil craft gains sp ...
, or
fin A fin is a thin component or appendage attached to a larger body or structure. Fins typically function as foils that produce lift or thrust, or provide the ability to steer or stabilize motion while traveling in water, air, or other fluids. Fin ...
s, are used to push against the water to create a normal force to provide thrust, propelling the animal through water. Sea turtles and penguins beat their paired hydrofoils to create lift. Some paired fins, such as pectoral fins on leopard sharks, can be angled at varying degrees to allow the animal to rise, fall, or maintain its level in the water column. The reduction of fin surface area helps to minimize drag, and therefore increase efficiency. Regardless of size of the animal, at any particular speed, maximum possible lift is proportional to (wing area) x (speed)2. Dolphins and whales have large, horizontal caudal hydrofoils, while many fish and sharks have vertical caudal hydrofoils. Porpoising (seen in cetaceans, penguins, and pinnipeds) may save energy if they are moving fast. Since drag increases with speed, the work required to swim unit distance is greater at higher speeds, but the work needed to jump unit distance is independent of speed. Seals propel themselves through the water with their caudal tail, while sea lions create thrust solely with their pectoral flippers.


Drag powered swimming

As with moving through any fluid, friction is created when molecules of the fluid collide with organism. The collision causes drag against moving fish, which is why many fish are streamlined in shape. Streamlined shapes work to reduce drag by orienting elongated objects parallel to the force of drag, therefore allowing the current to pass over and taper off the end of the fish. This streamlined shape allows for more efficient use of energy locomotion. Some flat-shaped fish can take advantage of pressure drag by having a flat bottom surface and curved top surface. The pressure drag created allows for the upward lift of the fish. Appendages of aquatic organisms propel them in two main and biomechanically extreme mechanisms. Some use lift powered swimming, which can be compared to flying as appendages flap like wings, and reduce drag on the surface of the appendage. Others use drag powered swimming, which can be compared to oars rowing a boat, with movement in a horizontal plane, or paddling, with movement in the parasagittal plane. Drag swimmers use a cyclic motion in which they push water back in a power stroke, and return their limb forward in the return or recovery stroke. When they push water directly backwards, this moves their body forward, but as they return their limbs to the starting position, they push water forward, which will thus pull them back to some degree, and so opposes the direction that the body is heading. This opposing force is called drag. The return-stroke drag causes drag swimmers to employ different strategies than lift swimmers. Reducing drag on the return stroke is essential for optimizing efficiency. For example, ducks paddle through the water spreading the webs of their feet as they move water back, and then when they return their feet to the front they pull their webs together to reduce the subsequent pull of water forward. The legs of water beetles have little hairs which spread out to catch and move water back in the power stroke, but lay flat as the appendage moves forward in the return stroke. Also, one side of a water beetle leg is wider than the others and is held perpendicular to the motion when pushing backward, but the leg rotates when the limb returns forward, so the thinner side catches less water.Lauder, G., and Jayne, B. Pectoral fin Locomotion in Fishes: Testing Drag-Based Models Using Three-Dimensional Kinematics. Oxford Journals. 2011. Drag swimmers experience a lessened efficiency in swimming due to resistance which affects their optimum speed. The less drag a fish experiences, the more it will be able to maintain higher speeds. Morphology of the fish can be designed to reduce drag, such as streamlining the body. The cost of transport is much higher for the drag swimmer, and when deviating from its optimum speed, the drag swimmer is energetically strained much more than the lift swimmer. There are natural processes in place to optimize energy use, and it is thought that adjustments of metabolic rates can compensate in part for mechanical disadvantages.Ohlberger, J., G. Staaks, and F. Hoker. Swimming efficiency and the influence of morphology on swimming costs in fishes. J Comp Physiol B. 176: 17-25. 2005. Semi-aquatic animals compared to fully aquatic animals exhibit exacerbation of drag. Design that allows them to function out of the water limits the efficiency possible to be reached when in the water. In water swimming at the surface exposes them to resistive wave drag and is associated with a higher cost than submerged swimming. Swimming below the surface exposes them to resistance due to return strokes and pressure, but primarily friction. Frictional drag is due to fluid viscosity and morphology characteristics. Pressure drag is due to the difference of water flow around the body and is also affected by body morphology. Semi-aquatic organisms encounter increased resistive forces when in or out of the water, as they are not specialized for either habitat. The morphology of otters and beavers, for example, must meet needs for both environments. Their fur decreases streamlining and creates additional drag. The platypus may be a good example of an intermediate between drag and lift swimmers because it has been shown to have a rowing mechanism which is similar to lift-based pectoral oscillation. The limbs of semi-aquatic organisms are reserved for use on land and using them in water not only increases the cost of locomotion, but limits them to drag-based modes.Fish, Frank E. Biomechanics and Energetics in Aquatic and Semiaquatic Mammals: Platypus to Whale. Chicago Journals. 2011. Although they are less efficient, drag swimmers are able to produce more thrust at low speeds than lift swimmers. They are also thought to be better for maneuverability due to the large thrust produced.Walker, J., and M. Westneat. Mechanical Performance of aquatic rowing and flying. The Royal Society. 10.1098. 2000.


Amphibians

Most of the Amphibia have a larval state, which has inherited anguilliform motion, and a laterally compressed tail to go with it, from fish ancestors. The corresponding tetrapod adult forms, even in the tail-retaining sub-class
Urodeles Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
, are sometimes aquatic to only a negligible extent (as in the genus
Salamandra ''Salamandra'' is a genus of seven species of salamanders localized in central and southern Europe, Northern Africa, and western Asia. List of species References External links Salamandraat Fauna Europaea * Salamandraat Animal Diversity W ...
, whose tail has lost its suitability for aquatic propulsion), but the majority of
Urodeles Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
, from the newts to the giant salamander Megalobatrachus, retain a laterally compressed tail for a life that is aquatic to a considerable degree, which can use in a carangiform motion.Lighthill , MJ. Hydromechanics of Aquatic Animal Propulsion- Annu. Rev. Fluid. Mech. - 1969 1-413-446. Of the tailless amphibians (the
frogs 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 ...
and
toads Toad is a common name for certain frogs, especially of the family Bufonidae, that are characterized by dry, leathery skin, short legs, and large bumps covering the parotoid glands. A distinction between frogs and toads is not made in scienti ...
of the sub-class Anura) the majority are aquatic to an insignificant extent in adult life, but in that considerable minority that are mainly aquatic we encounter for the first time the problem of adapting the tailless-tetrapod structure for aquatic propulsion. The mode that they use is unrelated to any used by fish. With their flexible back legs and webbed feet they execute something close to the leg movements of a human 'breast stroke,' rather more efficiently because the legs are better streamlined.


Reptiles

From the point of view of aquatic propulsion, the descent of modern members of the class
Reptilia Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
from archaic tailed
Amphibia Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbore ...
is most obvious in the case of the order
Crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
(
crocodiles Crocodiles (family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to include all extant memb ...
and
alligators An alligator is a large reptile in the Crocodilia order in the genus ''Alligator'' of the family Alligatoridae. The two extant species are the American alligator (''A. mississippiensis'') and the Chinese alligator (''A. sinensis''). Additionall ...
), which use their deep, laterally compressed tails in an essentially carangiform mode of propulsion (see Fish locomotion#Carangiform). Terrestrial
snakes Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more joi ...
, in spite of their 'bad' hydromechanical shape with roughly circular cross-section and gradual posterior taper, swim fairly readily when required, by an anguilliform propulsion (see Fish locomotion#Anguilliform).
Cheloniidae Cheloniidae is a family of typically large marine turtles that are characterised by their common traits such as, having a flat streamlined wide and rounded shell and almost paddle-like flippers for their forelimbs. They are the only sea turtles t ...
(true
turtles Turtles are an order (biology), order of reptiles known as Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) an ...
) have found a beautiful solution to the problem of
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
swimming through the development of their forelimbs into flippers of high-aspect-ratio wing shape, with which they imitate a bird's propulsive mode more accurately than do the eagle-rays themselves.


Fin and flipper locomotion

Aquatic reptiles such as sea turtles (see also
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked tu ...
s) and extinct species like
Pliosauroids Pliosauroidea is an extinct clade of plesiosaurs, known from the earliest Jurassic to early Late Cretaceous. They are best known for the subclade Thalassophonea, which contained crocodile-like short-necked forms with large heads and massive to ...
predominantly use their pectoral flippers to propel themselves through the water and their pelvic flippers for maneuvering. During swimming they move their pectoral flippers in a dorso-ventral motion, causing forward motion. During swimming, they rotate their front flippers to decrease drag through the water column and increase efficiency. Newly hatched sea turtles exhibit several behavioral skills that help orientate themselves towards the ocean as well as identifying the transition from sand to water. If rotated in the pitch, yaw or roll direction, the hatchlings are capable of counteracting the forces acting upon them by correcting with either their pectoral or pelvic flippers and redirecting themselves towards the open ocean. Among mammals
otariids An eared seal or otariid or otary is any member of the marine mammal family Otariidae, one of three groupings of pinnipeds. They comprise 15 extant species in seven genera (another species became extinct in the 1950s) and are commonly known eit ...
(
fur seals Fur seals are any of nine species of pinnipeds belonging to the subfamily Arctocephalinae in the family '' Otariidae''. They are much more closely related to sea lions than true seals, and share with them external ears (pinnae), relatively lon ...
) swim primarily with their front flippers, using the rear flippers for steering, and
phocids The earless seals, phocids or true seals are one of the three main groups of mammals within the seal lineage, Pinnipedia. All true seals are members of the family Phocidae (). They are sometimes called crawling seals to distinguish them from t ...
(
true seals The earless seals, phocids or true seals are one of the three main groups of mammals within the seal lineage, Pinnipedia. All true seals are members of the family Phocidae (). They are sometimes called crawling seals to distinguish them from th ...
) move the rear flippers laterally, pushing the animal through the water.


Escape reactions

Some arthropods, such as
lobster Lobsters are a family (biology), family (Nephropidae, Synonym (taxonomy), synonym Homaridae) of marine crustaceans. They have long bodies with muscular tails and live in crevices or burrows on the sea floor. Three of their five pairs of legs ...
s and
shrimp Shrimp are crustaceans (a form of shellfish) with elongated bodies and a primarily swimming mode of locomotion – most commonly Caridea and Dendrobranchiata of the decapod order, although some crustaceans outside of this order are refer ...
s, can propel themselves backwards quickly by flicking their tail, known as lobstering or the
caridoid escape reaction The caridoid escape reaction, also known as lobstering or tail-flipping, refers to an innate escape mechanism in marine and freshwater crustaceans such as lobsters, krill, shrimp and crayfish. The reaction, most extensively researched in crayfis ...
. Varieties of fish, such as teleosts, also use fast-starts to escape from predators. Fast-starts are characterized by the muscle contraction on one side of the fish twisting the fish into a C-shape. Afterwards, muscle contraction occurs on the opposite side to allow the fish to enter into a steady swimming state with waves of undulation traveling alongside the body. The power of the bending motion comes from fast-twitch muscle fibers located in the central region of the fish. The signal to perform this contraction comes from a set of Mauthner cells which simultaneously send a signal to the muscles on one side of the fish. Mauthner cells are activated when something startles the fish and can be activated by visual or sound-based stimuli. Fast-starts are split up into three stages. Stage one, which is called the preparatory stroke, is characterized by the initial bending to a C-shape with small delay caused by hydrodynamic resistance. Stage two, the propulsive stroke, involves the body bending rapidly to the other side, which may occur multiple times. Stage three, the rest phase, cause the fish to return to normal steady-state swimming and the body undulations begin to cease. Large muscles located closer to the central portion of the fish are stronger and generate more force than the muscles in the tail. This asymmetry in muscle composition causes body undulations that occur in Stage 3. Once the fast-start is completed, the position of the fish has been shown to have a certain level of unpredictability, which helps fish survive against predators. The rate at which the body can bend is limited by resistance contained in the inertia of each body part. However, this inertia assists the fish in creating propulsion as a result of the momentum created against the water. The forward propulsion created from C-starts, and steady-state swimming in general, is a result of the body of the fish pushing against the water. Waves of undulation create rearward momentum against the water providing the forward thrust required to push the fish forward.Wakeling, J.M. "Biomechanics of fast-start swimming in fish." Comparative Biochemistry and Physiology Part A.131 (2001): 31-40.


Efficiency

The Froude propulsion efficiency is defined as the ratio of power output to the power input: where U1 = free stream velocity and U2 = jet velocity. A good efficiency for carangiform propulsion is between 50 and 80%.


Minimizing drag

Pressure differences occur outside the boundary layer of swimming organisms due to disrupted flow around the body. The difference on the up- and down-stream surfaces of the body is pressure drag, which creates a downstream force on the object. Frictional drag, on the other hand, is a result of fluid viscosity in the
boundary layer In physics and fluid mechanics, a boundary layer is the thin layer of fluid in the immediate vicinity of a bounding surface formed by the fluid flowing along the surface. The fluid's interaction with the wall induces a no-slip boundary cond ...
. Higher
turbulence In fluid dynamics, turbulence or turbulent flow is fluid motion characterized by chaotic changes in pressure and flow velocity. It is in contrast to a laminar flow, which occurs when a fluid flows in parallel layers, with no disruption between ...
causes greater frictional drag.
Reynolds number In fluid mechanics, the Reynolds number () is a dimensionless quantity that helps predict fluid flow patterns in different situations by measuring the ratio between inertial and viscous forces. At low Reynolds numbers, flows tend to be domi ...
(Re) is the measure of the relationships between inertial and viscous forces in flow ((animal's length x animal's velocity)/kinematic viscosity of the fluid). Turbulent flow can be found at higher Re values, where the boundary layer separates and creates a wake, and laminar flow can be found at lower Re values, when the boundary layer separation is delayed, reducing wake and kinetic energy loss to opposing water momentum.Feldkamp, S.D. "Swimming in the California sea lion: morphometrics, drag, and energetics." J. exp. Biol. 131, 117-135 (1987). The body shape of a swimming organism affects the resulting drag. Long, slender bodies reduce pressure drag by streamlining, while short, round bodies reduce frictional drag; therefore, the optimal shape of an organism depends on its niche. Swimming organisms with a fusiform shape are likely to experience the greatest reduction in both pressure and frictional drag. Wing shape also affects the amount of drag experienced by an organism, as with different methods of stroke, recovery of the pre-stroke position results in the accumulation of drag. High-speed ram ventilation creates laminar flow of water from the gills along the body of an organism. The secretion of mucus along the organism's body surface, or the addition of long-chained polymers to the velocity gradient, can reduce frictional drag experienced by the organism.


Buoyancy

Many aquatic/marine organisms have developed organs to compensate for their weight and control their
buoyancy Buoyancy (), or upthrust, is an upward force exerted by a fluid that opposes the weight of a partially or fully immersed object. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus the p ...
in the water. These structures, make the density of their bodies very close to that of the surrounding water. Some hydrozoans, such as siphonophores, has gas-filled floats; the Nautilus, Sepia, and Spirula ( Cephalopods) have chambers of gas within their shells; and most
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tele ...
fish and many
lantern fish Lanternfishes (or myctophids, from the Greek μυκτήρ ''myktḗr'', "nose" and ''ophis'', "serpent") are small mesopelagic fish of the large family Myctophidae. One of two families in the order Myctophiformes, the Myctophidae are represente ...
(Myctophidae) are equipped with swim bladders. Many aquatic and marine organisms may also be composed of low-density materials. Deep-water teleosts, which do not have a swim bladder, have few lipids and proteins, deeply ossified bones, and watery tissues that maintain their buoyancy. Some
sharks Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimorp ...
' livers are composed of low-density lipids, such as hydrocarbon
squalene Squalene is an organic compound. It is a triterpenoid with the formula C30H50. It is a colourless oil, although impure samples appear yellow. It was originally obtained from shark liver oil (hence its name, as ''Squalus'' is a genus of sharks). A ...
or
wax esters A wax ester (WE) is an ester of a fatty acid and a fatty alcohol. Wax esters comprise the main components of three commercially important waxes: carnauba wax, candelilla wax, and beeswax.. Wax esters are formed by combining one fatty acid with ...
(also found in Myctophidae without swim bladders), which provide buoyancy. Swimming animals that are denser than water must generate lift or adapt a
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
lifestyle. Movement of the fish to generate hydrodynamic lift is necessary to prevent sinking. Often, their bodies act as
hydrofoils A hydrofoil is a lifting surface, or foil, that operates in water. They are similar in appearance and purpose to aerofoils used by aeroplanes. Boats that use hydrofoil technology are also simply termed hydrofoils. As a hydrofoil craft gains sp ...
, a task that is more effective in flat-bodied fish. At a small tilt angle, the lift is greater for flat fish than it is for fish with narrow bodies. Narrow-bodied fish use their fins as hydrofoils while their bodies remain horizontal. In sharks, the
heterocercal Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as s ...
tail shape drives water downward, creating a counteracting upward force while thrusting the shark forward. The lift generated is assisted by the pectoral fins and upward-angle body positioning. It is supposed that
tunas A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae (mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max length ...
primarily use their pectoral fins for lift. Buoyancy maintenance is metabolically expensive. Growing and sustaining a buoyancy organ, adjusting the composition of biological makeup, and exerting physical strain to stay in motion demands large amounts of energy. It is proposed that lift may be physically generated at a lower energy cost by swimming upward and gliding downward, in a "climb and glide" motion, rather than constant swimming on a plane.


Temperature

Temperature can also greatly affect the ability of aquatic organisms to move through water. This is because temperature not only affects the properties of the water, but also the organisms in the water, as most have an ideal range specific to their body and metabolic needs.
Q10 (temperature coefficient) The ''Q''10 temperature coefficient is a measure of temperature sensitivity based on the chemical reactions. The ''Q''10 is calculated as: : Q_=\left( \frac \right )^ where; : ''R'' is the rate : ''T'' is the temperature in Celsius degrees o ...
, the factor by which a rate increases at a 10 °C increase in temperature, is used to measure how organisms' performance relies on temperature. Most have increased rates as water becomes warmer, but some have limits to this and others find ways to alter such effects, such as by endothermy or earlier recruitment of faster muscle. For example, ''Crocodylus porosus'', or estuarine crocodiles, were found to increase swimming speed from 15 °C to 23 °C and then to have peak swimming speed from 23 °C to 33 °C. However, performance began to decline as temperature rose beyond that point, showing a limit to the range of temperatures at which this species could ideally perform.


Submergence

The more of the animal's body that is submerged while swimming, the less energy it uses. Swimming on the surface requires two to three times more energy than when completely submerged. This is because of the bow wave that is formed at the front when the animal is pushing the surface of the water when swimming, creating extra drag.


Secondary evolution

While
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s lost many of their natural adaptations to swimming when they evolved onto the land, many have re-evolved the ability to swim or have indeed returned to a completely aquatic lifestyle. Primarily or exclusively aquatic animals have re-evolved from terrestrial tetrapods multiple times: examples include amphibians such as
newts A newt is a salamander in the subfamily Pleurodelinae. The terrestrial juvenile phase is called an eft. Unlike other members of the family Salamandridae, newts are semiaquatic, alternating between aquatic and terrestrial habitats. Not all aqua ...
,
reptiles Reptiles, as most commonly defined are the animals in the Class (biology), class Reptilia ( ), a paraphyletic grouping comprising all sauropsid, sauropsids except birds. Living reptiles comprise turtles, crocodilians, Squamata, squamates (lizar ...
such as
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to inclu ...
s,
sea turtle Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerhead, ...
s, ichthyosaurs,
plesiosaurs The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared i ...
and
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on ...
s,
marine mammal Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their reli ...
s such as
whales Whales are a widely distributed and diverse group of fully aquatic placental marine mammals. As an informal and colloquial grouping, they correspond to large members of the infraorder Cetacea, i.e. all cetaceans apart from dolphins and ...
,
seals Seals may refer to: * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, or "true seal" ** Fur seal * Seal (emblem), a device to impress an emblem, used as a means of a ...
and
otter Otters are carnivorous mammals in the subfamily Lutrinae. The 13 extant otter species are all semiaquatic, aquatic, or marine, with diets based on fish and invertebrates. Lutrinae is a branch of the Mustelidae family, which also includes wea ...
s, and
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s such as
penguin Penguins (order (biology), order List of Sphenisciformes by population, Sphenisciformes , family (biology), family Spheniscidae ) are a group of Water bird, aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: on ...
s. Many species of
snakes Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more joi ...
are also aquatic and live their entire lives in the water. Among invertebrates, a number of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
species have adaptations for aquatic life and locomotion. Examples of aquatic insects include
dragonfly 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 ...
larvae, water boatmen, and
diving beetle The Dytiscidae – based on the Greek ''dytikos'' (δυτικός), "able to dive" – are the predaceous diving beetles, a Family (biology), family of water beetles. They occur in virtually any freshwater habitat around the world, but a ...
s. There are also aquatic spiders, although they tend to prefer other modes of locomotion under water than swimming proper. Even though primarily terrestrial tetrapods have lost many of their adaptations to swimming, the ability to swim has been preserved or re-developed in many of them. It may never have been completely lost. Examples are: Some breeds of
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. Do ...
swim recreationally. Umbra, a world record-holding dog, can swim 4 miles (6.4 km) in 73 minutes, placing her in the top 25% in human long-distance swimming competitions. The
fishing cat The fishing cat (''Prionailurus viverrinus'') is a medium-sized wild cat of South and Southeast Asia. Since 2016, it is listed as Vulnerable on the IUCN Red List. Fishing cat populations are threatened by destruction of wetlands and have declin ...
is one wild species of cat that has evolved special adaptations for an aquatic or semi-aquatic lifestyle – webbed digits. Tigers and some individual jaguars are the only big cats known to go into water readily, though other big cats, including lions, have been observed swimming. A few domestic cat breeds also like swimming, such as the
Turkish Van The Turkish Van (pronounced Von) is a naturally occurring breed of domestic cat that originated in the Lake Van area of modern-day Turkey, part of Armenian Highlands. The Van is classified as a semi-long hair, but it has two lengths of hair, ...
. Horses,
moose The moose (in North America) or elk (in Eurasia) (''Alces alces'') is a member of the New World deer subfamily and is the only species in the genus ''Alces''. It is the largest and heaviest extant species in the deer family. Most adult ma ...
, and
elk The elk (''Cervus canadensis''), also known as the wapiti, is one of the largest species within the deer family, Cervidae, and one of the largest terrestrial mammals in its native range of North America and Central and East Asia. The common ...
are very powerful swimmers, and can travel long distances in the water. Elephants are also capable of swimming, even in deep waters. Eyewitnesses have confirmed that
camel A camel (from: la, camelus and grc-gre, κάμηλος (''kamēlos'') from Hebrew or Phoenician: גָמָל ''gāmāl''.) is an even-toed ungulate in the genus ''Camelus'' that bears distinctive fatty deposits known as "humps" on its back. C ...
s, including
dromedary The dromedary (''Camelus dromedarius'' or ;), also known as the dromedary camel, Arabian camel, or one-humped camel, is a large even-toed ungulate, of the genus ''Camelus'', with one hump on its back. It is the tallest of the three species of ...
and Bactrian camels, can swim, despite the fact that there is little deep water in their natural habitats. Both domestic and wild
rabbit Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit speci ...
s can swim. Domestic rabbits are sometimes trained to swim as a circus attraction. A wild rabbit famously swam in an apparent attack on U.S. President
Jimmy Carter James Earl Carter Jr. (born October 1, 1924) is an American politician who served as the 39th president of the United States from 1977 to 1981. A member of the Democratic Party (United States), Democratic Party, he previously served as th ...
's boat when it was threatened in its natural habitat. The
guinea pig The guinea pig or domestic guinea pig (''Cavia porcellus''), also known as the cavy or domestic cavy (), is a species of rodent belonging to the genus ''Cavia'' in the family Caviidae. Breeders tend to use the word ''cavy'' to describe the ani ...
(or cavy) is noted as having an excellent swimming ability.
Mice A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
can swim quite well. They do panic when placed in water, but many lab mice are used in the
Morris water maze The Morris water navigation task, also known as the Morris water maze (not to be confused with '' water maze''), is a behavioral procedure mostly used with rodents. It is widely used in behavioral neuroscience to study spatial learning and me ...
, a test to measure learning. When mice swim, they use their tails like
flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
and kick with their legs. Many snakes are excellent swimmers as well. Large adult
anaconda Anacondas or water boas are a group of large snakes of the genus '' Eunectes''. They are found in tropical South America. Four species are currently recognized. Description Although the name applies to a group of snakes, it is often used ...
s spend the majority of their time in the water, and have difficulty moving on land. Many monkeys can naturally swim and some, like the proboscis monkey,
crab-eating macaque The crab-eating macaque (''Macaca fascicularis''), also known as the long-tailed macaque and referred to as the cynomolgus monkey in laboratories, is a cercopithecine primate native to Southeast Asia. A species of macaque, the crab-eating macaqu ...
, and
rhesus macaque The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of Old World monkey. There are between six and nine recognised subspecies that are split between two groups, the Chinese-derived and the Indian-derived. Generally ...
swim regularly.


Human swimming

Swimming has been known amongst humans since prehistoric times; the earliest record of swimming dates back to
Stone Age The Stone Age was a broad prehistoric period during which stone was widely used to make tools with an edge, a point, or a percussion surface. The period lasted for roughly 3.4 million years, and ended between 4,000 BC and 2,000 BC, with t ...
paintings from around 7,000 years ago. Competitive swimming started in
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
around 1800 and was part of the first modern
1896 Summer Olympics The 1896 Summer Olympics ( el, Θερινοί Ολυμπιακοί Αγώνες 1896, Therinoí Olympiakoí Agónes 1896), officially known as the Games of the I Olympiad ( el, Αγώνες της 1ης Ολυμπιάδας, Agónes tis 1is Ol ...
in
Athens Athens ( ; el, Αθήνα, Athína ; grc, Ἀθῆναι, Athênai (pl.) ) is both the capital and largest city of Greece. With a population close to four million, it is also the seventh largest city in the European Union. Athens dominates ...
, though not in a form comparable to the contemporary events. It was not until 1908 that regulations were implemented by the
International Swimming Federation FINA (french: Fédération internationale de natation, en, International Swimming Federation, link=yes) (to be renamed as World Aquatics by ) is the international federation recognised by the International Olympic Committee (IOC) for administer ...
to produce
competitive swimming Swimming is an individual or team racing sport that requires the use of one's entire body to move through water. The sport takes place in pools or open water (e.g., in a sea or lake). Competitive swimming is one of the most popular Olympic ...
.


See also

*
Animal locomotion Animal locomotion, in ethology, is any of a variety of methods that animal (biology), animals use to move from one place to another. Some modes of locomotion are (initially) self-propelled, e.g., running, swimming, jumping, flying, hopping, soari ...
* Aquatic *
Fish fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as see ...
*
Locomotion in space Locomotion in space includes all actions or methods used to move one's body in microgravity conditions through the outer space environment. Locomotion in these conditions is different from locomotion in a gravitational field. There are many facto ...
*
Robot locomotion Robot locomotion is the collective name for the various methods that robots use to transport themselves from place to place. Wheeled robots are typically quite energy efficient and simple to control. However, other forms of locomotion may be more ...
* Role of skin in locomotion *
Terrestrial locomotion Terrestrial locomotion has evolved as animals adapted from aquatic to terrestrial environments. Locomotion on land raises different problems than that in water, with reduced friction being replaced by the increased effects of gravity. As viewed ...
* Tradeoffs for locomotion in air and water *
Undulatory locomotion Undulatory locomotion is the type of motion characterized by wave-like movement patterns that act to propel an animal forward. Examples of this type of gait include crawling in snakes, or swimming in the lamprey. Although this is typically the ...


References

{{DEFAULTSORT:Aquatic Locomotion Ethology Swimming Animal locomotion