Information about Mobbing Behavior

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The Great Tit, a passerine bird, employs both mobbing behavior and alarm calls.
In behavioral ecology, mobbing behavior is an antipredator behavior which occurs when individuals of a certain species mob a predator by cooperatively attacking or harassing it, usually in order to protect their offspring. A simple definition of mobbing is an assemblage of individuals around a potentially dangerous predator.<ref name="Dominey" /> This is most frequently seen in avian species, though it is also known to occur in other social animals. While mobbing has evolved independently in many species, it only tends to be present in those whose young are frequently preyed on. This behavior may complement cryptic adaptations in the offspring themselves, such as camouflage and hiding. Mobbing calls may be used to summon nearby individuals to cooperate in the attack.

Mobbing in birds

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Nesting Kittiwakes.
Birds that breed in colonies such as gulls are widely seen to attack intruders, including encroaching humans.[1] Behavior includes flying about the intruder, dive bombing, loud squawking and defecating on the predator. Costs of mobbing behavior include the risk of engaging with predators, as well as energy expended in the process. Black-headed Gulls are one species which aggressively engages intruding predators, such as Carrion Crows. Classic experiments on this species by Hans Kruuk involved placing hen eggs at intervals from a nesting colony, and recording the percentage of successful predation events as well as the probability of the crow being subjected to mobbing.[2] The results showed decreasing mobbing with increased distance from the nest, which was correlated with increased predation success. Mobbing may function by reducing the predator's ability to locate nests, in other words as a distraction, since predators cannot focus on locating eggs while they are under direct attack.

Adaptationist hypotheses regarding why an organism should engage in such risky behavior have been suggested by Eberhard Curio,[3] including advertising their physical fitness and hence uncatchability (much like stotting behavior in gazelles), distracting predators from finding their offspring, warning their offspring, luring the predator away, allowing offspring to learn to recognize the predator species,[4] directly injuring the predator or attracting a predator of the predator itself. The much lower frequency of attacks between nesting seasons suggests such behavior may have evolved due to its benefit for the mobber's young. Niko Tinbergen argued that the mobbing was a source of confusion to gull chick predators, distracting them from searching for prey .[5] Indeed, an intruding carrion crow can only avoid incoming attacks by facing its attackers, which prevents it from locating its target.<ref name="Alcock" />

Besides experimental research, the comparative method can also be employed to investigate hypotheses such as those given by Curio above. For example, closely related species such as the Kittiwake do not show mobbing behavior. The kittiwake's cliff nests are almost completely inaccessible to possible predators due to gusty winds and the shear nature of the cliffs they nest in, meaning its young are not at risk to predation like the Black-headed Gull.[6] This is an example of an evolutionary pattern known as divergent evolution.

Mobbing is thought to carry risks to roosting predators, including suffering harm from the mobbing birds or the risk of attracting larger, more dangerous predators. Birds at risk of mobbing such as owls have adapted cryptic plumage and hidden roosting sites in order to reduce this danger.[7]

In other animals

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The occurrence of mobbing behavior across widely different taxa, including California Ground Squirrels, is evidence of convergent evolution.
Another way the comparative method can be used here is by comparing gulls with distantly related organisms. This approach relies on the existence of convergent evolution, where distantly related organisms evolve the same trait due to similar selection pressures. As mentioned, many bird species such as the swallows also show mobbing of predators, however even more distantly related species including mammals are known to engage in this behavior. One example is California Ground Squirrels, which are known to distract predators such as the rattlesnake and gopher snake from locating their nest burrows by kicking sand into their eyes.[8] This social species also uses alarm calls.

Mobbing has also been observed in fishes, for example bluegills have been seen to attack snapping turtles.[9] Bluegills, which form large nesting colonies, were seen to attack both released and naturally occurring turtles, which may function to advertise their presence, drive the predator from the area, or aid in cultural transmission of predator recognition.

Mobbing calls

Mobbing calls are signals made by the mobbing species while harassing a predator. These differ from alarm calls, which allow con-specifics to escape from the predator. The Great Tit, a European songbird uses such a signal to call on nearby birds to harass a perched bird of prey, such as an owl. This call occurs in the 4.5kHz range,<ref name="Alcock" /> and is effective in traveling long distances. However, when their prey are in flight, they employ an alarm signal in the 7-8kHz range. This call is less effective at travelling great distances, but is much more difficult for both owls and hawks to hear (and detect the direction from which the call came).[10] In the case of the alarm call, it is disadvantageous to the sender if the predator picks up on the signal, hence selection has favored those birds able to hear and employ calls in this higher frequency range.

Mobbing calls may also be part of an animal's arsenal in harassing the predator - for example studies of Phainopepla mobbing calls indicate it may serve to enhance the swooping attack on the predators, including Scrub Jays. In this species the mobbing call is smoothly upsweeping, and is made when swooping down in an arc beside the predator. This call was also heard during agonistic interactions with conspecifics, and may serve additionally or alternatively as an alarm call to their mate.[11]

Evolution

The evolution of mobbing behaviour is explained using evolutionary stable strategies which are in turn based on Game Theory.[12]

Mobbing involves risks (costs) to the individual and benefits (payoffs) to the individual and others. The individuals themselves are often genetically related and it is increasingly studied with the Gene-centered view of evolution by considering inclusive fitness (the carrying on of one's genes through one's family members), rather than merely benefit to the individual.

By cooperating to successfully drive away predators all individuals involved increase their chances of survival and reproduction. An individual stands little chance against a larger predator, but when a large group is involved, the risk to each group member is minimized. By being in a large group, the risk for a particular individual is reduced or diluted. This so-called dilution effect proposed by W. D. Hamilton is another way of explaining the benefits of cooperation by selfish individuals. Lanchester's laws also provide an insight into the advantages of attacking in a large group rather than individually. [13][14]

Another interpretation involves the use of the handicap principle. Here the idea is that a mobbing bird, by apparently putting itself at risk, displays its status and health so as to be preferred by potential partners.[15]

References

1. ^ Alcock, John (1998). Animal Behavior: An Evolutionary Approach, 6th edition, Sunderland: Sinauer Associates. ISBN 0-87893-009-4. 
2. ^ Kruuk, H. (1964). Predators and anti-predator behaviour of the black-headed gull Larus ridibundus, Behaviour Supplements (11). Leiden: E.J. Brill. OCLC 1502972. 
3. ^ Curio, E. (1978). "The adaptive significance of avian mobbing. I. Teleonomic hypotheses and predictions". Zeitschrift fur Tierpsychologie 48: 175-183. 
4. ^ Curio, E.; U. Ernst; W. Vieth (1978). "Cultural Transmission of Enemy Recognition: One Function of Mobbing". Science 202: 899-901. 
5. ^ Tinbergen, Niko [1960] (1967). The herring gull's world: a study of the social behavior of birds. New York: Lyons and Bulford. ISBN 1-55821-049-0. 
6. ^ Cullen, E. (1957). "Adaptations in the kittiwake to cliff nesting". Ibis 99: 275-302. 
7. ^ Ditte K. Hendrichsen, Peter Christiansen, Elsemarie K. Nielsen, Torben Dabelsteen & Peter Sunde, (2006) "Exposure affects the risk of an owl being mobbed – experimental evidence" Journal of Avian Biology 37(1): 13–18
8. ^ Coss, Richard G. (1997). "Individual Variation in the Antisnake Behavior of California Ground Squirrels (Spermophilus beecheyi)". Journal of Mammalogy 78 (2): 294-310. 
9. ^ Dominey, Wallace J. (1983). "Mobbing in Colonially Nesting Fishes, Especially the Bluegill, Lepomis macrochirus". Copeia 1983 (4): 1086-1088. DOI:10.2307/1445113. 
10. ^ Brown, C. H. (1982). "Ventriloquial and locatable vocalizations in birds". Zeitschrift fur Tierpsychologies 59: 338-350. 
11. ^ Leger, Daniel W.; Laura F. Carroll (1981). "Mobbing Calls of the Phainopepla". The Condor 83 (4): 377-380. DOI:10.2307/1367509. Retrieved on 2007-06-12. 
12. ^ Parker, Geoffrey A., Manfred Milinski (1997). "Cooperation under Predation Risk: A Data-Based ESS Analysis.". Proceedings: Biological Sciences 264 (1385): 1239-1247. 
13. ^ Kelly, Kevin (1994). Out of control: the new biology of machines, social systems and the economic world. Boston: Addison-Wesley. ISBN 0-201-48340-8. 
14. ^ Hamilton, W. D. 1971. Geometry for the selfish herd. J. theor. Biol. 31:295-311.
15. ^ Arnold, K. E. (2000). "Group Mobbing Behaviour and Nest Defence in a Cooperatively Breeding Australian Bird". Ethology 106: 385-393. 

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Behavioral ecology is the study of the ecological and evolutionary basis for animal behavior, and the roles of behavior in enabling an animal to adapt to its environment (both intrinsic and extrinsic).
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Antipredator adaptations are adaptations developed over evolutionary time, which assist prey organisms in their constant struggle against their predators. There are several ways antipredator adaptations can be classified, such as behavioral or non-behavioral or by taxonomic groups.
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Behavior or behaviour
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Mobbing is a new term referring to a group behavioural phenomenon in workplaces and a type of animal behaviour. In a different sense, it is a criminal offence in Scotland.

Mobbing

In the book MOBBING: Emotional Abuse in the American Workplace
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predation describes a biological interaction where a predator organism feeds on another living organism or organisms known as prey.[1] Predators may or may not kill their prey prior to feeding on them.
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offspring are the product of reproduction, a new organism produced by one or more parents.

Collective offspring may be known as a brood or progeny in a more general way.
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Aves
Linnaeus, 1758

Orders

About two dozen - see section below

Birds (class Aves) are bipedal, warm-blooded, egg-laying vertebrate animals.
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crypsis is the ability of an organism to avoid observation. A cryptic animal may do this through camouflage, nocturnality, subterranean lifestyle, transparency[1], or mimicry.
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Camouflage, also known as cryptic coloration or concealing coloration, allows an otherwise visible organism or object to remain indiscernible from the surrounding environment. Examples include a tiger's stripes and the battledress of a modern soldier.
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Concealment or hiding is obscuring something from view or rendering it inconspicuous, the opposite of exposure. A military term is CCD: camouflage, concealment and deception (looks the same as the surroundings, can not be seen, looks like something else, respectively); in a
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Co-operation or co-operative behaviours are terms used to describe behaviours by biological organisms which are beneficial to other members of the same species. There are several competing theories which help to explain why natural selection favours some types of
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Laridae
Vigors, 1825

Genera

Larus
Rissa
Pagophila
Rhodostethia
Xema
Creagus

Gulls are birds in the family Laridae.
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Defecation is the act or process by which organisms eliminate solid or semisolid waste material (feces) from the digestive tract via the anus. Humans expel feces anywhere from a few times daily to a few times weekly; sloths can go a week without expelling.
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L. ridibundus

Binomial name
Larus ridibundus
Linnaeus, 1766, European seas

The Black-headed Gull (Larus ridibundus
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Adaptationism is a set of methods in the evolutionary sciences for distinguishing the products of adaptation from traits that arise through other processes. It is employed in fields such as ethology and evolutionary psychology that are concerned with identifying adaptations.
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A scientific hypothesis is a hypothesis (a testable conjecture) which is used as a tentative explanation of an observation, but which has not yet been fully tested by the prediction validation process for a scientific theory.
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Stotting or pronking is a gait of quadrupeds, particularly gazelles (e.g. Thomson's Gazelles), involving jumping high into the air. This may occur during pursuit by a predator, typically a cheetah or lion. It might also occur during play.
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Animal communication is any behaviour on the part of one animal that has an effect on the current or future behaviour of another animal. The study of animal communication, sometimes called zoosemiotics
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kin selection.

The concept was formalized by JBS Haldane (1955)[1] and W. D. Hamilton (1963)[2], while the actual term "kin selection" may first have been coined by John Maynard Smith (1964)[3]
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Niko Tinbergen

Nikolaas "Niko" Tinbergen
(1907-1988)
Born March 15 1907(1907--)
The Hague, Netherlands
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Rissa
Stephens, 1826

Species

Rissa tridactyla
Rissa brevirostris

The Kittiwakes (genus Rissa) are two closely related seabird species in the gull family Laridae.
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