Animal locomotion on the surface layer of water is the study of

animal locomotion In ethology, animal locomotion is any of a variety of methods that animals use to move from one place to another. Some modes of locomotion are (initially) self-propelled, e.g., running, swimming, jumping, flight, flying, hopping, soaring and gli ...

in the case of small

animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...

s that live on the

surface A surface, as the term is most generally used, is the outermost or uppermost layer of a physical object or space. It is the portion or region of the object that can first be perceived by an observer using the senses of sight and touch, and is ...

layer of

water Water is an inorganic compound with the chemical formula . It is a transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance. It is the main constituent of Earth's hydrosphere and the fluids of all known liv ...

, relying on

surface tension Surface tension is the tendency of liquid surfaces at rest to shrink into the minimum surface area possible. Surface tension (physics), tension is what allows objects with a higher density than water such as razor blades and insects (e.g. Ge ...

to stay afloat. There are two types of animal locomotion on water, determined by the ratio of the animal's

weight In science and engineering, the weight of an object is a quantity associated with the gravitational force exerted on the object by other objects in its environment, although there is some variation and debate as to the exact definition. Some sta ...

to the water's

: those whose weight is supported by the surface tension at rest, and can therefore easily remain on the water's surface without much exertion, and those whose weight is not supported by the water's surface tension at rest, and must therefore exert additional motion in a direction parallel to the water's surface in order to remain above it. A creature such as the

basilisk lizard ''Basiliscus'' is a genus of large corytophanid lizards, commonly known as basilisks, which are endemic to southern Mexico, Central America, and northern South America. The genus contains four species, which are commonly known as the Jesus Chr ...

, often dubbed the 'Jesus lizard', has a weight which is larger than the surface tension can support, and is widely known for running across the surface of water. Another example, the

western grebe The western grebe (''Aechmophorus occidentalis'') is a species in the grebe family of water birds. Folk names include "dabchick", "swan grebe" and "swan-necked grebe". Western grebe fossils from the Late Pleistocene of southwest North America we ...

, performs a mating ritual that includes running across the surface of water. Surface living animals such as the

water strider The Gerridae are a family of insects in the order Hemiptera, commonly known as water striders, water skeeters, water scooters, water bugs, pond skaters, water skippers, water gliders, water skimmers or puddle flies. They are true bugs of the ...

typically have

hydrophobic In chemistry, hydrophobicity is the chemical property of a molecule (called a hydrophobe) that is seemingly repelled from a mass of water. In contrast, hydrophiles are attracted to water. Hydrophobic molecules tend to be nonpolar and, thu ...

feet covered in small

hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and ...

s that prevent the feet from breaking the surface and becoming wet. Another insect known to walk on the water surface is the ant species ''

Polyrhachis sokolova '' Polyrhachis sokolova'' is a species of ant from Australia and New Guinea that recently was discovered to be capable of surviving tidal inundations. Researchers at the James Cook University gained attention in 2006 after finding and reportin ...

''. The pygmy gecko ('' Coleodactylus amazonicus''), due to its small size and hydrophobic skin, is also able to walk on the water surface. According to

biophysicist Biophysics is an interdisciplinary science that applies approaches and methods traditionally used in physics to study biological phenomena. Biophysics covers all scales of biological organization, from molecular to organismic and populations ...

David L. Hu, there are at least 342 species of water striders. As striders increase in size, their legs become proportionately longer, with ''

Gigantometra gigas ''Gigantometra'' is a monotypic genus of water-strider bugs, containing the species ''Gigantometra gigas'': its name indicating that it is the largest species in its family Gerridae. It has been found in pools of fast-flowing, subtropical and t ...

'' having a length of over 20 cm requiring a surface tension force of about 40 millinewtons. Water striders generate thrust by shedding

vortices In fluid dynamics, a vortex (: vortices or vortexes) is a region in a fluid in which the flow revolves around an axis line, which may be straight or curved. Vortices form in stirred fluids, and may be observed in smoke rings, whirlpools in th ...

in the water: a series of U-shaped vortex filaments is created during the power stroke. The two free ends of the "U" are attached to the water surface. These vortices transfer enough (backward) momentum to the water to propel the animal forwards (note that some momentum is transferred by capillary waves; see Denny's paradox for a more detailed discussion.)

Meniscus climbing

To pass from the water surface to land, a water-walking insect must contend with the slope of the

meniscus Meniscus may refer to: *Meniscus (anatomy), crescent-shaped fibrocartilaginous structure that partly divides a joint cavity *Meniscus (liquid) In physics (particularly liquid statics), the meniscus (: menisci, ) is the curve in the upper sur ...

at the water's edge. Many such insects are unable to climb this meniscus using their usual propulsion mechanism. David Hu and coworker John W. M. Bush have shown that such insects climb meniscuses by assuming a fixed body posture. This deforms the water surface and generates capillary forces that propels the insect up the slope without moving its appendages. Hu and Bush conclude that meniscus climbing is an unusual means of propulsion in that the insect propels itself in a quasi-static configuration, without moving its appendages. Biolocomotion is generally characterized by the transfer of muscular strain energy to the kinetic and

gravitational potential energy Gravitational energy or gravitational potential energy is the potential energy an object with mass has due to the gravitational potential of its position in a gravitational field. Mathematically, it is the minimum Work (physics), mechanical work t ...

of the creature, and the kinetic energy of the suspending fluid. In contrast, meniscus climbing has a different energy pathway: by deforming the free surface, the insect converts muscular strain to the surface energy that powers its ascent.

Marangoni propulsion

Many insects, including some terrestrial insects, can release a

surfactant Surfactants are chemical compounds that decrease the surface tension or interfacial tension between two liquids, a liquid and a gas, or a liquid and a solid. The word ''surfactant'' is a Blend word, blend of "surface-active agent", coined in ...

and propel themselves using the

Marangoni effect The Marangoni effect (also called the Gibbs–Marangoni effect) is the mass transfer along an Interface (chemistry), interface between two phases due to a gradient of the surface tension. In the case of temperature dependence, this phenomenon may ...

. Hu and Bush report that '' Microvelia'' can attain a peak speed of 17 cm/s, which is twice its peak walking speed, using Marangoni propulsion. Marangoni propulsion by a

wetting Wetting is the ability of a liquid to displace gas to maintain contact with a solid surface, resulting from intermolecular interactions when the two are brought together. These interactions occur in the presence of either a gaseous phase or ...

arthropod is precisely analogous to a soap boat but the situation for insects such as water striders is more complex. Hu and Bush state that "for nonwetting arthropods, the transfer of chemical to kinetic energy is more subtle, as the Marangoni stress must be communicated across the creature’s complex surface layer".

Sailing

Velella ''Velella'' is a monospecific genus of hydrozoa in the family Porpitidae. Its only known species is ''Velella velella'', a cosmopolitan (widely distributed) free-floating hydrozoan that lives on the surface of the open ocean. It is commonly kn ...

'', the by-the-wind sailor, is a

cnidaria Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...

n with no means of propulsion other than

sailing Sailing employs the wind—acting on sails, wingsails or kites—to propel a craft on the surface of the ''water'' (sailing ship, sailboat, raft, Windsurfing, windsurfer, or Kitesurfing, kitesurfer), on ''ice'' (iceboat) or on ''land'' (Land sa ...

. A small rigid sail projects into the air and catches the wind. ''Velella'' sails always align along the direction of the wind where the sail may act as an

aerofoil An airfoil (American English) or aerofoil (British English) is a streamlined body that is capable of generating significantly more lift than drag. Wings, sails and propeller blades are examples of airfoils. Foils of similar function designed ...

, so that the animals tend to sail downwind at a small angle to the wind.

References

* John W. M. Bush and David L. Hu
''WALKING ON WATER: Biolocomotion at the Interface''
Annual Review of Fluid Mechanics, vol. 38, p. 339-369, January 2006.

External links

(photos) {{locomotion Animal locomotion Water transport

Meniscus climbing

Marangoni propulsion

Sailing

See also

References

External links