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Stotting (also called pronking or pronging) is a behavior of quadrupeds, particularly
gazelle A gazelle is one of many antelope species in the genus ''Gazella'' . This article also deals with the seven species included in two further genera, '' Eudorcas'' and '' Nanger'', which were formerly considered subgenera of ''Gazella''. A third ...
s, in which they spring into the air, lifting all four feet off the ground simultaneously. Usually, the legs are held in a relatively stiff position. Many explanations of stotting have been proposed, though for several of them there is little evidence either for or against. The question of why prey animals stot has been investigated by evolutionary biologists including
John Maynard Smith John Maynard Smith (6 January 1920 – 19 April 2004) was a British theoretical and mathematical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics un ...
, C. D. Fitzgibbon, and
Tim Caro Timothy M. Caro (born 1951) is a British evolutionary ecologist known for his work on conservation biology, animal behaviour, anti-predator defences in animals, and the function of zebra stripes. He is the author of several textbooks on these subj ...
; all of them conclude that the most likely explanation given the available evidence is that it is an honest signal to
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
s that the stotting animal would be difficult to catch. Such a signal is called "honest" as it is not deceptive in any way, and would benefit both predator and prey; the predator as it avoids a costly and unproductive chase, and the prey as it does not get chased.


Etymology

''Stot'' is a common Scots and Northern English verb meaning "bounce" or "walk with a bounce". Uses in this sense include stotting a ball off a wall, and rain stotting off a pavement. Pronking comes from the
Afrikaans Afrikaans (, ) is a West Germanic language that evolved in the Dutch Cape Colony from the Dutch vernacular of Holland proper (i.e., the Hollandic dialect) used by Dutch, French, and German settlers and their enslaved people. Afrikaans gr ...
verb ''pronk-'', which means "show off" or "strut", and is a
cognate In historical linguistics, cognates or lexical cognates are sets of words in different languages that have been inherited in direct descent from an etymological ancestor in a common parent language. Because language change can have radical ef ...
of the English verb "prance".


Taxonomic distribution

Stotting occurs in several deer species of
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
, including
mule deer The mule deer (''Odocoileus hemionus'') is a deer indigenous to western North America; it is named for its ears, which are large like those of the mule. Two subspecies of mule deer are grouped into the black-tailed deer. Unlike the related whi ...
,
pronghorn The pronghorn (, ) (''Antilocapra americana'') is a species of artiodactyl (even-toed, hoofed) mammal indigenous to interior western and central North America. Though not an antelope, it is known colloquially in North America as the American a ...
, and
Columbian black-tailed deer Two forms of black-tailed deer or blacktail deer that occupy coastal woodlands in the Pacific Northwest of North America are subspecies of the mule deer (''Odocoileus hemionus''). They have sometimes been treated as a species, but virtually all r ...
, when a
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
is particularly threatening, and in a variety of
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, ...
species from
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
, including
Thomson's gazelle Thomson's gazelle (''Eudorcas thomsonii'') is one of the best known species of gazelles. It is named after explorer Joseph Thomson and is sometimes referred to as a "tommie". It is considered by some to be a subspecies of the red-fronted gazell ...
and
springbok The springbok (''Antidorcas marsupialis'') is a medium-sized antelope found mainly in south and southwest Africa. The sole member of the genus ''Antidorcas'', this bovid was first described by the German zoologist Eberhard August Wilhelm ...
. It is also said to occur in the
blackbuck The blackbuck (''Antilope cervicapra''), also known as the Indian antelope, is an antelope native to India and Nepal. It inhabits grassy plains and lightly forested areas with perennial water sources. It stands up to high at the shoulder. Ma ...
, a species found in India. Stotting occurs in domesticated
livestock Livestock are the domesticated animals raised in an agricultural setting to provide labor and produce diversified products for consumption such as meat, eggs, milk, fur, leather, and wool. The term is sometimes used to refer solely to ani ...
such as
sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to domesticate ...
and
goats The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of th ...
, typically only in young animals.


Possible explanations

Stotting makes a prey animal more visible, and uses up time and energy that could be spent on escaping from the predator. Since it is dangerous, the continued performance of stotting by prey animals must bring some benefit to the animal (or its family group) performing the behavior. Several possible explanations have been proposed, namely that stotting may be: # A good means of rapid escape or jumping over obstructions. However, this cannot be true in Thomson's gazelles because these prey animals do not stot when a predator is less than approximately 40 m away. # An anti-ambush behavior; animals living in tall grass may leap into the air to detect potential predators. There is some evidence for this. # An alarm signal to other members of the herd that a predator is hazardously close thereby increasing the survival rate of the herd. This would be an instance of
group selection Group selection is a proposed mechanism of evolution in which natural selection acts at the level of the group, instead of at the level of the individual or gene. Early authors such as V. C. Wynne-Edwards and Konrad Lorenz argued that the behavi ...
, a theory heavily criticised by evolutionary biologists. # A socially cohesive behavior to escape predators by coordinated stotting, thereby making it more difficult for a predator to target any individual during an attack. This too would be group selection, subject to the same objections. # An honest signal of the animal's fitness. Stotting could be a way of deterring pursuit by warning a predator of the animal's unsuitability as prey: the prey benefits by not being chased (because it is in fact very fit); the predator benefits by not wasting time chasing an animal it is unlikely to catch. This signalling explanation avoids the
group selection Group selection is a proposed mechanism of evolution in which natural selection acts at the level of the group, instead of at the level of the individual or gene. Early authors such as V. C. Wynne-Edwards and Konrad Lorenz argued that the behavi ...
connotations of the "alarm signal" and "socially cohesive" escape hypotheses. # An instance of
Amotz Zahavi Amotz Zahavi ( he, אמוץ זהבי) (August 14, 1928 – May 12, 2017) was an Israeli evolutionary biologist, a Professor in the Department of Zoology at Tel Aviv University, and one of the founders of the Society for the Protection of Natur ...
's
handicap principle The handicap principle is a hypothesis proposed by the biologist Amotz Zahavi to explain how evolution may lead to "honest" or reliable signalling between animals which have an obvious motivation to bluff or deceive each other. It suggests that ...
, whereby stotting is signalling to predators that the animal is so fit it can escape even if it deliberately slows itself down with some apparently useless behavior (i.e. stotting). # A predator detection signal whereby the animal signals to the predator that it has been seen and therefore does not have the advantage of surprise. Many such signals exist in different groups of animals. Again, this would be an honest pursuit deterrence signal, benefiting the prey by not being chased (because it can be seen to be aware of the predator and ready to escape immediately) and benefitting the predator by not wasting time stalking prey when it has already been seen. Evidence for this hypothesis is that
cheetah The cheetah (''Acinonyx jubatus'') is a large cat native to Africa and central Iran. It is the fastest land animal, estimated to be capable of running at with the fastest reliably recorded speeds being , and as such has evolved specialized ...
s abandon more hunts when their gazelle prey stots, and when they do give chase to a stotting gazelle, they are far less likely to make a kill. However, gazelles stot less often to cheetahs (which stalk and would therefore probably give up when detected) than to
African wild dog The African wild dog (''Lycaon pictus''), also called the painted dog or Cape hunting dog, is a wild canine which is a native species to sub-Saharan Africa. It is the largest wild canine in Africa, and the only extant member of the genus '' Lyca ...
s, which "
course Course may refer to: Directions or navigation * Course (navigation), the path of travel * Course (orienteering), a series of control points visited by orienteers during a competition, marked with red/white flags in the terrain, and corresponding ...
" (chase prey relentlessly, not relying on surprise). # A fitness display to potential mates in a
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...
process rather than an
antipredator adaptation Anti-predator adaptations are mechanisms developed through evolution that assist prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, namely by avo ...
., South African Lodges # Play, especially in young animals, which may help to prepare them for adult life. In favor of this hypothesis, stotting is sometimes observed in immature animals; against this is the fact that stotting is generally seen in adult prey responding to predators. The English evolutionary biologist
John Maynard Smith John Maynard Smith (6 January 1920 – 19 April 2004) was a British theoretical and mathematical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics un ...
concludes that "the natural explanation is that stotting is an index of condition and of escape capability", used as a signal especially to coursing predators. He also observes that "it is hard to see how it could be a handicap", unless perhaps it is a signal to other gazelles of the same species. C. D. Fitzgibbon agrees that it is most likely an honest signal of condition.
Tim Caro Timothy M. Caro (born 1951) is a British evolutionary ecologist known for his work on conservation biology, animal behaviour, anti-predator defences in animals, and the function of zebra stripes. He is the author of several textbooks on these subj ...
comments that there is insufficient evidence "to support or refute a startle effect of stotting, prey signalling its health, the Social Cohesion hypothesis or nalarm function of stotting"; in his view, stotting informs a predator that it has been detected. File:Look Ma, I can Jump!.jpg, A lamb stotting File:Impala-stotting.jpg, An adult male
black-faced impala The impala or rooibok (''Aepyceros melampus'') is a medium-sized antelope found in eastern and southern Africa. The only extant member of the genus ''Aepyceros'' and tribe Aepycerotini, it was first described to European audiences by German ...
stotting in
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
File:Stotting gazelle.jpg, An
impala The impala or rooibok (''Aepyceros melampus'') is a medium-sized antelope found in eastern and southern Africa. The only extant member of the genus ''Aepyceros'' and tribe Aepycerotini, it was first described to European audiences by Ger ...
stotting


See also

*
Aposematism Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste ...
*
Deimatic behaviour Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey anima ...
*
Signalling theory Within evolutionary biology, signalling theory is a body of theoretical work examining communication between individuals, both within species and across species. The central question is when organisms with conflicting interests, such as in se ...


References


External links

{{Ethology Ethology Terrestrial locomotion