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Stotting

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Overview

Stotting (also pronking or pronging) is a behavior of quadrupeds, particularly gazelles, 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 and the back may be arched with the head pointing downward. Many explanations of stotting have been proposed; there is evidence that at least in some cases it is an honest signal to predators that the stotting animal is not worth pursuing.

Etymology

Stot is a common Scots and Geordie verb meaning "bounce" or "walk with a bounce."[1] Uses in this sense include stotting a ball off a wall, and rain stotting off a pavement. Pronking comes from the Afrikaans verb pronk-, which means "show off" or "strut", and is a cognate of the English verb "prance".[2]

Occurrence

Stotting occurs in several deer species of North America, including mule deer and pronghorn,[3] Columbian black-tailed deer when a predator is particularly threatening,[4] and in a variety of ungulate species from Africa, including Thomson's Gazelle and Springbok.[5]

Stotting occurs in domesticated livestock such as sheep, but is typically performed only by young animals.[6]

Possible explanations

Stotting makes a prey animal more visible[7] and uses up energy and time that could be spent escaping. Therefore, it must bring some benefit to the animal performing the behavior. A number of possible explanations have been proposed for stotting.[8][9] Stotting may be:

  1. 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.[5][10]
  2. An anti-ambush behavior; animals living in tall grass may leap into the air to detect potential predators.[8]
  3. An alarm signal to other members of the herd that a predator is hazardously close thereby increasing the survival rate of the herd.[a][8]
  4. 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 (much like the suggestion that zebra stripes cause motion dazzle).[8]
  5. 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 theory explanation avoids the group selection connotations of the "alarm signal" and "socially cohesive" escape hypotheses.[8][10]
  6. An instance of Amotz Zahavi's handicap principle, 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), just as the best horse in a handicap race is the one carrying the heaviest saddlebags.[11]
  7. 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 cheetahs 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.[9] However, gazelles stot less often to cheetahs (which stalk and would therefore probably give up when detected) than to African wild dogs, which "course" (chase prey relentlessly, not relying on surprise).[10]
  8. A fitness display to potential mates in a sexual selection process rather than an antipredator adaptation.[12]
  9. 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, however against the play hypothesis is that stotting is "generally" seen in adult prey responding to predators.[10]

The English evolutionary biologist John Maynard Smith 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.[5]

See also

Notes

  1. ^ This would be an instance of kin selection or group selection.

References

The Handicap Principle: A Missing Piece of Darwin's Puzzle
Amotz Zahavi:::Avishag Zahavi (1997)
Ever since Darwin, animal behavior has intrigued and perplexed human observers. The elaborate mating rituals, lavish decorative displays, complex songs, calls, dances and many other forms of animal signaling raise fascinating questions. To what degree can animals communicate within their own species and even between species? What evolutionary purpose do such communications serve? Perhaps most importantly, what can animal signaling tell us about our own non-verbal forms of communication? In The Handicap Principle, Amotz and Ashivag Zahavi offer a unifying theory that brilliantly explains many previously baffling aspects of animal signaling and holds up a mirror in which ordinary human behaviors take on surprising new significance. The wide-ranging implications of the Zahavis' new theory make it arguably the most important advance in animal behavior in decades. Based on 20 years of painstaking observation, the Handicap Principle illuminates an astonishing variety of signaling behaviors in animals ranging from ants and ameba to peacocks and gazelles. Essentially, the theory asserts that for animal signals to be effective they must be reliable, and to be reliable they must impose a cost, or handicap, on the signaler. When a gazelle sights a wolf, for instance, and jumps high into the air several times before fleeing, it is signaling, in a reliable way, that it is in tip-top condition, easily able to outrun the wolf. (A human parallel occurs in children's games of tag, where faster children will often taunt their pursuer before running). By momentarily handicapping itself--expending precious time and energy in this display--the gazelle underscores the truthfulness of its signal. Such signaling, the authors suggest, serves the interests of both predator and prey, sparing each the exhaustion of a pointless chase. Similarly, the enormous cost a peacock incurs by carrying its elaborate and weighty tail-feathers, which interfere with food gathering, reliably communicates its value as a mate able to provide for its offspring. Perhaps the book's most important application of the Handicap Principle is to the evolutionary enigma of animal altruism. The authors convincingly demonstrate that when an animal acts altruistically, it handicaps itself--assumes a risk or endures a sacrifice--not primarily to benefit its kin or social group but to increase its own prestige within the group and thus signal its status as a partner or rival. Finally, the Zahavis' show how many forms of non-verbal communication among humans can also be explained by the Handicap Principle. Indeed, the authors suggest that non-verbal signals--tones of voice, facial expressions, body postures--are quite often more reliable indicators of our intentions than is language. Elegantly written, exhaustively researched, and consistently enlivened by equal measures of insight and example, The Handicap Principle illuminates virtually every kind of animal communication. It not only allows us to hear what animals are saying to each other--and to understand why they are saying it--but also to see the enormously important role non-verbal behavior plays in human communication.
  1. ^ Definition of stot
  2. ^ Definition of pronk
  3. ^ Roosevelt, Theodore (1905). Outdoor Pastimes of an American Hunter. C. Scribner's Sons. 
  4. ^ Stankowich, Theodore; Coss, Richard (2007). "Effects of Risk Assessment, Predator Behavior, and Habitat on Escape Behavior in Columbian Black-Tailed Deer". Behavioral Ecology 18 (2): 358–367. doi:10.1093/beheco/arl086. 
  5. ^ a b c Maynard Smith, John; Harper, David (2003). Animal Signals. Oxford University Press. pp. 61–63 [1]. 
  6. ^ Simmons, Paula; Carol Ekarius (2001). Storey's Guide to Raising Sheep. North Adams, MA: Storey Publishing LLC. ISBN 978-1-58017-262-2. 
  7. ^ Anon (19 June 1986). "How the cheetah lost its stotts". New Scientist: 34. 
  8. ^ a b c d e Alcock, J. (2009). Animal Behavior. (Ninth ed.). Massachusetts: Sinauer Associates Inc.
  9. ^ a b Caro, TM (1986). "The functions of stotting in Thomson’s gazelles: Some tests of the predictions". Animal Behaviour 34: 663–684. doi:10.1016/S0003-3472(86)80052-5. 
  10. ^ a b c d FitzGibbon, CD; Fanshawe, JH (August 1988). "Stotting in Thomson’s gazelles: an honest signal of condition". Behavioral Ecology and Sociobiology 23 (2): 69–74 [2]. doi:10.1016/S0003-3472(86)80052-5. 
  11. ^ Zahavi, Amotz (1997). The handicap principle: a missing piece of Darwin's puzzle. Oxford: Oxford University Press. ISBN 0-19-510035-2. 
  12. ^ Herbivores of the Pilanesberg National Park I, South African Lodges
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