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(New page: {{CompPsy}} [[Silver gulls will often mob predators who approach their nesting site.<ref name="Alcock">Alcock, J. (1998) ''Animal Behavior: An Ev...)
 
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[[Image:Red legged gulls.jpg|right|thumb|[[Silver gull]]s will often mob predators who approach their nesting site.<ref name="Alcock">Alcock, J. (1998) ''Animal Behavior: An Evolutionary Approach'' (8th edition). Sinauer Associates, Inc. Sunderland, Massachusetts. ISBN 0-87893-009-4</ref>]]
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{{CompPsy}}
'''Antipredator adaptations''' are [[adaptation]]s developed over [[evolution]]ary time, which assist [[prey]] organisms in their constant [[evolutionary arms race|struggle]] against their [[predator]]s. There are several ways antipredator adaptations can be classified, such as [[behavior]]al or non-behavioral or by [[taxonomic]] groups. <!-- In this article these adaptations will be classified by dividing predation into four stages<ref name="Alcock" />: detection, attack, capture and consumption. At each stage adaptations that maximize a prey organisms chances of survival have evolved, which in turn drives responding adaptions in their predators. This interaction over long periods of time is known as [[co-evolution]].
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[[Image:Canis lupus pack surrounding Bison.jpg|thumb|A lone [[bison]] is standing its ground against a pack of [[wolf|wolves]], thereby increasing its chance of survival]]
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'''Antipredator adaptations''' are [[evolution]]ary [[adaptation]]s developed over time, which assist [[prey]] organisms in their constant [[evolutionary arms race|struggle]] against their [[predator]]s.
   
==Avoiding detection==
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A predator's acquisition of a [[food source]] can be divided into four stages: detection, attack, capture and consumption.<ref name="Alcock">{{cite book |author=John Alcock |title=Animal Behavior: An Evolutionary Approach |edition=8th |year=1998 |publisher=Sinauer |isbn=0-87893-009-4 }}</ref><ref>Endler (1991) In ''Behavioural Ecology'', 3rd ed. (Krebs & Davies), pp. 169–196.</ref> At every stage in this predatory sequence, adaptations that maximize the prey's chance of survival have evolved.
For a predator to locate a potential meal, it must first identify a suitable prey organism. Prey, however, have many adaptive characteristics which make such a task difficult.
 
   
===Crypsis===
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== Animal adaptations ==
{{Main|Crypsis}} -->
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[[File:Orange oak leaf bottom.JPG|thumb|250px|right|The butterfly ''[[Kallima inachus]]'' masquerades as a dead leaf to avoid detection]]
'''Antipredator behaviors''' range from [[chameleon]]s and [[octopus]]es that change color in order to better camouflage themselves, to [[crab]]s and [[Geometrid Moth|geometrid moth caterpillars]] that decorate their bodies to help them hide, to [[Batesian mimicry]] in insects, to [[Thomson's Gazelle|Thomson’s gazelles]] that [[stot]] so as to advertise unprofitability to predators, to [[selfish herd]] acts performed by [[Adelie Penguin|adelie penguins]] when they attempt to enter water inhabited with [[leopard seals]]. Their behaviors are extraordinarily varied from species to species and some of them are quite odd; some prey actually fight back.
 
   
Three extremely odd defense behaviors are found in the [[Sea Cucumber|sea cuke]] (cucumber), the ''[[Camponotus saundersi]]'' (Malaysian ant) and the [[horned lizard]] found in the [[Southwestern United States|desert southwest]] and [[Mexico]]. The sea cuke has a unique way of fighting off danger. Like other [[echinoderms]] the cuke has a type of [[collagen]] in its skin capable of excreting or absorbing more water effectively changing from a “liquid” to a “solid.” They can turn their bodies into mush, climb through small cracks and then solidify into small lumps so that they cannot be extracted. Their more desperate response is called evisceration. The cuke effectively turns itself inside out by excreting portions of its digestive tract and, on occasion, other organs like the respiratory tree or gonads. The cuke does this because its [[Cuvierian tubule|Cuvierian tubules]], located in its hindgut, contain toxic chemicals. The toxins can actually kill other fish. Any predator that comes into contact with this noxious substance is likely to think twice about consuming the cuke. The cuke can then regenerate its digestive tract and continue to survive.
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=== Avoiding detection ===
   
The horned lizard found in the desert southwest of the U.S. has an odd defense mechanism as well. When threatened the lizard increases pressure in its sinus cavities until the blood vessels in the corners of its eyes burst, squirting blood at the attacker.
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==== Avoiding being seen ====
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{{main|Camouflage}}
   
''Camponotus saundersi'', an [[ant]] species found in [[Malaysia]], also has a very interesting defense. The colony is divided up into different functional groups, one of which is soldier ants. These soldiers are charged with defending the colony at all costs. If battle ensues, these ants will actually self-destruct. They have two large glands that run the entire length of their body, and when they become stressed the ant contracts its abdominal muscles causing the glands to explode, spraying poison in all directions.
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For a predator to locate a potential meal, it must first [[Prey detection|identify an organism as prey]]. Prey, however, have many adaptive characteristics which make such a task difficult. [[Crypsis]] is the ability of an animal, predator or prey, to make itself hard to see.<ref>{{cite book | title=Dazzled and Deceived: Mimicry and Camouflage | publisher=Yale | author=Forbes, Peter | year=2009 | pages=50-51}}</ref>
These three examples seem to be some of the more bizarre ways that animals defend themselves against predation.
 
   
==References==
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[[Camouflage]] is one heavily utilised method, and involves the mimicking of the colour and patterns of the environment. This is achieved by external [[biological pigment|pigmentation]] patterns, and resting immobile and silent on an appropriate substrate, as do [[Egyptian Nightjar]]. Many marine animals use [[countershading]] or [[Camouflage#Crypsis_by_counterillumination|counterillumination]] to reduce their visibility from above and below. Some, such as the [[Mimic Octopus]], can rapidly change their pattern and colour to provide a better defense. Some animals provide structures on their bodies for plant life to grow, camouflaging its host. Other animals [[Camouflage#Camouflage_by_mimesis|masquerade]] as specific, inedible objects: the [[Celaenia excavata|Bird Dropping Spider]] and the [[Phobaeticus|''Phobaeticus'' Stick insect]]s are notable examples.<ref>[[Cott, H.B.]] (1940) ''Adaptive Coloration in Animals''. Methuen, London. (Stick insects: 334-335. Bird dropping spider, 330-332.)</ref>
<references />
 
[[Category:Predation]]
 
   
:fr:Moyen de défense
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[[File:Polymorphism in Cepaea nemoralis.jpg|thumb|center|500px|Polymorphism in the [[Grove snail]] is partly a response to predation by [[Song Thrush]]es]]
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Visual [[Polymorphism (biology)|Polymorphism]], the existence of different forms within a single species, is a strategy used to reduce predation risk. Predators make use of search images to differentiate edible objects from all other objects; this causes frequency-dependent "[[apostatic selection]]" in which morphs with rarer colours and patterns are less likely to be killed. The striking polymorphism of the [[Grove snail]] may be partly caused by apostatic selection by predators including the [[Song Thrush]].<ref name="CainSheppard">{{cite journal | url=http://www.genetics.org/content/39/1/89.full.pdf | title=Natural Selection in Cepaea | author=Cain, A.J. and Sheppard, P.M. | journal=Genetics | year=1954 | volume=39 | issue=1 | pages=89–116|pmid=17247470 | pmc=1209639}}</ref> There is evidence that polymorphic insect prey suffer less predation than single-morph species at a particular density, and can maintain higher population densities for a given rate of predation.<ref>Fullick & Greenwood (1979) Am. Nat. 113, 762-765.</ref><ref>{{aut|Edmunds, Malcolm.}} ''The Evolution of Cryptic Colour''. in ''Insect defenses: adaptive mechanisms and strategies of prey and predators'' (1990) Eds. David L. Evans, Justin O. Schmidt, pg 11.</ref>{{rp|11}}. This effect is more pronounced in tropical areas than temperate ones due to more intense and consistent interactions among different organisms.<ref>Masaki Hoso, Michio Hori
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The American Naturalist, Vol. 172, No. 5 (Nov., 2008), pp. 726-732</ref>
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==== Avoiding predators in space and time ====
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Animals alter the period in which they are awake in order to avoid predators.<ref name=Rosier>Rosier and Langkilde, 2011</ref> Generally, animals are either [[Diurnality|diurnal]] (active during the day), [[Nocturnality|nocturnal]] (active during the night), or [[crepuscular]] (active during twilight) depending on food availability and predator prevalence. For example, ''[[Dipodomys merriami]]'' kangaroo rats become crepuscular instead of nocturnal on full moon nights, when predators could see them more easily.<ref name=Rosier/>
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''[[Anolis sagrei]]'' lizards are more [[arboreal]] (living in trees) on islands where ground-living predators exist.<ref name=Rosier/>
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Highly mobile creatures such as seabirds [[Animal migration|migrate]] to avoid predators during the period in which they are most vulnerable, the [[breeding season]], for example moving to offshore [[island]]s to establish [[bird colony|colonies]] far from the reach of land predators.
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=== Avoiding attack ===
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{{further2|[[Aposematism]]|[[Mimicry]]}}
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Having been detected by a predator, many animals attempt to signal to a predator that they are not worth eating. Some animals make use of [[aposematic]] signals, for example bright [[warning colouration]], or sounds and smells, so as to advertise that they are poisonous.<ref>{{cite web |author=Juan Carlos Santos, Luis A. Coloma, David C. Cannatella |title=Multiple, recurring origins of aposematism and diet specialization in poison frogs |url=http://www.pnas.org/cgi/content/abstract/100/22/12792 |publisher=National Academy of Sciences |date=28 October 2003 |accessdate=2008-12-22 }}</ref> These patterns are often [[Convergent evolution|convergent]], with red or yellow coupled with black being widely recognised as dangerous. [[Batesian mimicry]] is the imitation by a harmless species of the warning signals of a harmful species directed at a common predator, as can be seen in a [[hoverfly]] mimicking a [[wasp]].<ref>[[Cott, H.B.]] (1940) ''Adaptive Coloration in Animals''. Methuen, London. pages 396-413.</ref>
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[[File:Automeris ioPCCP20040706-5706AB.jpg|thumb|The eyespots of ''[[Automeris io]]'']]
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Some animals, particularly [[gazelle]]s, are known to [[stot]], which, among other things, may advertise their unprofitability to predators.<ref>{{cite journal |author=Caro, T. M. |year=1986 |title=The functions of stotting in Thomson's gazelles: Some tests of the predictions. |journal=Animal Behaviour |issue=34 |pages=663–684 }}</ref> The moth ''[[Automeris io]]'' possesses [[Eyespot (mimicry)|eyespot]]s hidden on its hindwings. When under threat, the moth suddenly reveals these spots, aiming to startle its predator. Fearful of their own consumption, predators often retreat when startled. Eyespots are also used by some animals to trick a predator into thinking it has been spotted (when in fact, it may not have been), and so is less likely to succeed in pursuit.<ref>{{cite journal |doi=10.1017/S1464793105006810 |author=Stevens, Martin |title=The role of eyespots as anti-predator mechanisms, principally demonstrated in the Lepidoptera |journal=Biological Reviews |volume=80 |issue=4 |year=2005 |pages=573–588 |pmid=16221330 }}</ref>
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=== Avoiding capture ===
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Many animals have highly developed senses of [[Visual perception|sight]], [[Olfaction|smell]], and [[Hearing (sense)|hearing]] so that they can detect danger and escape. By frequently scanning and monitoring their surroundings, especially when in the open, prey can avoid attack by hoping to see a predator before it reaches the 'critical distance' for an attack. This is a standard defense mechanism for animals in open [[grasslands]] and [[prairies]]. It is also common for [[arboreal]] animals to scan both the ground around them for [[terrestrial animal|terrestrial]] predators, and the sky for aerial predators. Smaller animals may not venture too far from cover in burrows or the undergrowth, where they can quickly hide when danger approaches. [[Flying and gliding animals|Flight]] is of huge importance in the avoidance of predators in those species that possess it.<ref>{{cite web | url=http://www.nature.com/scitable/knowledge/library/behavior-under-risk-how-animals-avoid-becoming-23646978 | title=Behavior Under Risk: How Animals Avoid Becoming Dinner | publisher=Nature | date=2011 | accessdate=December 09, 2012 | author=Rosier, Renee L; Langkilde, Tracy}}</ref>
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=== Predator Recognition ===
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Learned Predator Recognition is influenced by pre-learning growth rate. That is the conclusion that was reached from the research done by Maud Ferrari, Grant Brown, Gary Bortolotti and Douglas Chivers on whether information learned about predators was influenced by prey growth trajectory before and after learning.<ref>{{cite journal|last=Ferrari|first=Maud C. O.|coauthors=Grant E. Brown • Gary R. Bortolotti • Douglas P. Chivers|title=Prey behaviour across antipredator adaptation types: how does growth trajectory influence learning of predators?|journal=Animal Cognition|year=2011|month=November|pages=809–816}}</ref>
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==== Safety in numbers ====
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Animals that are the frequent target of predation often make use of 'safety in numbers'. This results in a situation where any one herd member is unlikely to be preyed upon, and in high populations, [[predator satiation]] is likely to occur. [[Grazing]] mammals often feed in [[social group]]s, also known as [[herd]]s. Working as a group, a stalking predator is likely to be detected earlier, and when it attacks, the herd scatters, causing difficulty for the predator and allowing most, if not all, of the prey animals to escape. Prey animals may use [[alarm signal]]s to alert other herd members when a predator is sighted or sensed. Animals usually have a [[breeding season]], where all the members of the species spawn at the same time, in order to maximise their young's chance of survival. This is particularly pronounced in insects such as ''[[Magicicada]]'' and [[Mayfly|mayflies]], where millions of individuals emerge from pupation on the same day.<ref>{{cite web |url=http://insects.ummz.lsa.umich.edu/fauna/ |title=Periodical Cicada |author=John Cooley & Dave Marshall |date=9 January 2000 |publisher=University of Michigan |accessdate=2008-12-22 }}</ref>
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==== Fighting off predators ====
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[[File:MuskOxen.jpg|thumb|450px|[[Muskox]]en in defensive formation, horns ready, and highly alert]]
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Many animals use horns, claws, and teeth to fight off predators. Some can inject venom and toxins, and [[skunk]]s and [[bombardier beetle]]s<ref>{{cite book |author=Piper, Ross |authorlink=Ross Piper |title=Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals |publisher=[[Greenwood Press (publisher)|Greenwood Press]] |year=2007 |isbn=0-313-33922-8 }}</ref> spray noxious chemicals to deter attackers. [[Animal mobbing behaviour|Mobbing]], the harassing of a predator by many prey animals, is common in birds, and is usually done to protect the young in social colonies. The [[Apis cerana|Eastern Honeybee]] mobs invading [[hornet]]s, vibrating their flight muscles in order to raise the temperature around the hornet scout to lethal levels, rather than allowing the scout to bring others to their [[beehive]].<ref>{{cite journal |author=Masato Ono, Takeshi Igarashi, Eishi Ohno, and Masami Sasaki |date=28 September 1995 |title=Unusual thermal defence by a honeybee against mass attack by hornets |journal=Nature |issue=377 |pages=334–336 |url=http://www.nature.com/nature/journal/v377/n6547/abs/377334a0.html |accessdate=2008-12-22 | doi= 10.1038/377334a0 |volume=377}}</ref>
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==== Confusing the predator ====
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[[File:Zebras Serengeti.JPG|thumb|300px|A single [[zebra]] is hard to catch amongst a herd]]
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Some animals have evolved [[Camouflage#Dazzle_patterning|dazzle camouflage]], whereby instead of attempting to conceal themselves, they are patterned to cause motion dazzle, confusing a predator during an attack, and making it harder to select and track a target. An example is [[zebra]]s, which stand out in the [[savannah]] when stationary, but when moving rapidly ''en masse'', their stripes create a confusing, flickering mass in the eye of a predator such as a [[lion]].<ref name="Stevens Motion Dazzle">{{cite web | url=http://www.biomedcentral.com/1741-7007/9/81/abstract | title=BMC Biology: Motion dazzle | publisher=BMC Biology | pages=9:81 | doi=10.1186/1741-7007-9-81 | work=Motion dazzle and camouflage as distinct anti-predator defenses | date=25 November 2011 | accessdate=January 5, 2012 | author=Martin Stevens, William TL Searle, Jenny E Seymour, Kate LA Marshall, Graeme D Ruxton}}</ref>
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=== Avoiding consumption ===
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[[File:Fishing spider autotomy.jpg|thumb|This [[fishing spider]] has shed two of its legs to escape a predator]]
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Having been captured, an animal must prevent the predator from killing and eating it. Mechanical defenses in those that possess them, such as armour and spines prevent access to softer edible parts. [[Distraction display]]s are used to direct the attention of the predator away from some vital area, such as the head, or a nest of chicks.<ref name=ruxton>Ruxton, Graeme D; Thomas N. Sherratt; Michael Patrick Speed. (2004) Avoiding attack: the evolutionary ecology of crypsis, warning signals and mimicry. Oxford University Press. ISBN 0-19-852859-0. p. 198</ref> These can be visual, as in misdirecting eyespots, behavioural, such as a mother-bird feigning injury, or chemical, as seen in the production of [[cephalopod ink|ink clouds]] by squid and octopuses.
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Some animals (e.g. [[possum]]s) freeze in cover or [[apparent death|play dead]] when seen. Often this is accompanied by foul smells, as if their corpse is in an advanced state of decomposition, one which many animals would avoid consuming.<ref name=Pasteur>Pasteur, G. (1982). "[http://www.annualreviews.org/doi/pdf/10.1146/annurev.es.13.110182.001125 A classificatory review of mimicry systems]". ''Annual Review of Ecology and Systematics'' '''13''': 169–199.</ref>
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Some animals take a more drastic approach to defending themselves. [[Autotomy]], the shedding of a non-vital bodypart, is utilised by some species to escape the grasp of a predator. Many [[lizard]]s shed their tails when clasped, and arthropods will readily give up several legs if it allows their escape. When under threat, [[Holothuroidea|sea cucumber]]s rapidly [[Evisceration (autotomy)|eviscerate]], ejecting part or all of their digestive tract. This is done to either anchor the cucumber into a rock fissure, or to eject toxins at the predator.<ref>{{cite journal |title=Biomechanics of adhesion in sea cucumber cuvierian tubules (echinodermata, holothuroidea) |journal=Integrative and Comparative Biology |date=2002-12-01 |author=Patrick Flammang, Jerome Ribesse, Michel Jangoux |url=http://findarticles.com/p/articles/mi_qa4054/is_200212/ai_n9160035/pg_1 |accessdate=2008-12-22 | archiveurl= http://web.archive.org/web/20081226045104/http://findarticles.com/p/articles/mi_qa4054/is_200212/ai_n9160035/pg_1| archivedate= 26 December 2008}}</ref> [[Horned lizard]]s, when threatened, increase the pressure in their sinus cavities until the blood vessels in the corners of the eyes burst, squirting blood at the attacker.<ref>{{cite web |url=http://www.digimorph.org/resources/horned.phtml |title=About Horned Lizards |accessdate=2008-12-22 |author=Dr. Wendy Hodges |publisher=DigiMorph | archiveurl= http://web.archive.org/web/20090125170740/http://digimorph.org/resources/horned.phtml| archivedate= 25 January 2009 | deadurl= no}}</ref> [[Armoured cricket]]s and many other insect species also use this method, known as [[autohaemorrhaging]].<ref>[http://blogs.discovermagazine.com/discoblog/2009/07/28/see-it-to-believe-it-animals-vomit-and-spurt-blood-to-thwart-predators/ "See It to Believe It: Animals Vomit, Spurt Blood to Thwart Predators"], Allison Bond, ''[[Discover Magazine]]'' blog, 28 July 2009, retrieved 17 March 2010</ref> These animals are able to survive the loss of these [[Tissue (biology)|tissues]], and later [[Regeneration (biology)|regenerate]] them.
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The soldier ant caste of the [[Malaysia]]n species ''[[Camponotus saundersi]]'' undergoes a process known as [[autothysis]] to defend their [[ant colony]]. The soldier ants have two large glands that run the entire length of their body, and when stressed during battle, abdominal muscles contract, causing the glands to explode, killing the ant, but spraying poison in all directions.<ref>[http://www.springerlink.com/content/v0kjl37610tk2426/ Maschwitz, U. and E. Maschwitz, 1974. Platzende Arbeiterinnen: Eine neue Art der Feindabwehr bei sozialen Hautflüglern. Oecologia Berlin 14:289–294] (in German)</ref><ref name=bordereau>{{cite journal |author=C. Bordereau, A. Robert, V. Van Tuyen & A. Peppuy |year=1997 |title=Suicidal defensive behavior by frontal gland dehiscence in ''Globitermes sulphureus'' Haviland soldiers (Isoptera) |journal=[[Insectes Sociaux]] |volume=44 |issue=3 |pages=289–297 |doi=10.1007/s000400050049}}</ref>
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== Plant adaptations ==
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[[File:Aphid-sap.jpg|right|thumb|[[Aphid]] feeding on [[plant sap]]]]
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{{Main|Plant defense against herbivory}}
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Many plant species have, over the course of their evolutionary history, developed physical and chemical<ref>{{Cite web|url=http://www2.mcdaniel.edu/Biology/botf99/herbnew/aprodbc.htm |title=Biochemical defenses: secondary metabolites: |accessdate=2007-05-21 |work=Plant Defense Systems & Medicinal Botany}}</ref> defense mechanisms to deter [[herbivore]]s. [[Thorns, spines, and prickles]] are examples of physical mechanisms. [[Stinging nettle]]s for instance are covered in hollow hairs that can inject [[Irritation|irritant]] chemicals. [[Acacia]]s and [[rose]]s are among the many plants protected by spines and thorns. Prickles and spines are not effective against small insects such as [[aphid]]s however, so some plants, such as some of the [[Solanaceae]], supplement a bristly surface with sticky secretions that trap and kill small pests. Other plants such as [[holly]] have thick tough leaves covered with slippery way, making feeding difficult for small herbivores.<ref>{{Cite journal|author=Fernandes GW|year=1994|title=Plant mechanical defenses against insect herbivory|journal=Revista Brasileira de Entomologia|volume=38|issue=2|pages=421–433 [http://www.icb.ufmg.br/big/leeb/publicacoes/1994.Fernandes.pdf]}}</ref>
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Many ''[[Erica]]''s have a sticky perianth, and some ''[[Moraea]]''s have sticky [[Peduncle (botany)|peduncles]] that deter ants from plundering the flowers' [[nectar]] without contributing to their [[pollination]]. Many plants deter their enemies with repellent tastes or irritating or dangerous poisons in [[sap]] or in [[latex]].<ref name = vWvHvO>{{cite book | author = van Wyk, Ben-Erik; van Heerden, Fanie; van Oudtshoorn, Bosch| title = Poisonous Plants of South Africa | publisher = Briza | location = Pretoria | year = 2002 | isbn = 978-1-875093-30-4}}</ref> The herbivores in turn developed wide ranges of [[Herbivore adaptations to plant defense|counter-adaptations]].
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==See also==
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* [[Ultrasound avoidance]]
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* [[Predator avoidance in schooling fish]]
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== References ==
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{{reflist|2}}
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== Further reading ==
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* Caro, T. 2005. ''Antipredator Defenses in Birds and Mammals''. Chicago : University of Chicago Press. 591 pp.&nbsp;ISBN 0-226-09435-9 (hardcover version)
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* [[Cott, H.B.]] 1940. ''Adaptive Coloration in Animals''. London: Methuen.
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* Edmunds, M. 1974. ''Defence in Animals: A Survey of Anti-Predator Defences''. Harlow, Essex & NY: Longman ISBN 0-582-44132-3
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* Gabrielsen, G.W. & Smith, E.N.: ''Physiological responses associated with feigned death in the American Opossum''. Acta Phys. Scand. 123: 393-398. 1985.
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* Gabrielsen, G.W. & Smith, E.N.: ''Physiological responses to disturbance in animals''. In; Wildlife and Recreationists (R. Knight and K. Utzwiller. Island Press, Washington, D.C. pp.&nbsp;137–153. 1995.
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* Rosier, R.L. & Langkilde, T.: ''Behavior Under Risk: How Animals Avoid Becoming Dinner''. Nature Education Knowledge 2(11):8. 2011. [http://www.nature.com/scitable/knowledge/library/behavior-under-risk-how-animals-avoid-becoming-23646978 Full text]
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* Ruxton, G. D.; Speed, M. P.; Sherratt, T. N. (2004). ''Avoiding Attack. The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry''. Oxford: Oxford University Press. ISBN 0-19-852860-4
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* Steen, J.B., Gabrielsen, G.W. & Kanwischer, J.W.: ''Physiological aspects of freezing behavior in Willow Ptarmigan hens''. Acta Phys. Scand. 134: 299-304. 1988.
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{{Mimicry}}
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{{feeding}}
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{{modelling ecosystems|state=expanded}}
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[[Category:Antipredator adaptations|*]]
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[[Category:Evolutionary biology]]
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[[Category:Warning coloration]]
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Revision as of 15:31, December 21, 2012

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Animals · Animal ethology · Comparative psychology · Animal models · Outline · Index


Assessment | Biopsychology | Comparative | Cognitive | Developmental | Language | Individual differences | Personality | Philosophy | Social |
Methods | Statistics | Clinical | Educational | Industrial | Professional items | World psychology |

Animals · Animal ethology · Comparative psychology · Animal models · Outline · Index


File:Canis lupus pack surrounding Bison.jpg
A lone bison is standing its ground against a pack of wolves, thereby increasing its chance of survival

Antipredator adaptations are evolutionary adaptations developed over time, which assist prey organisms in their constant struggle against their predators.

A predator's acquisition of a food source can be divided into four stages: detection, attack, capture and consumption.[1][2] At every stage in this predatory sequence, adaptations that maximize the prey's chance of survival have evolved.

Animal adaptations

Orange oak leaf bottom
The butterfly Kallima inachus masquerades as a dead leaf to avoid detection
PhloxBotAdded by PhloxBot

Avoiding detection

Avoiding being seen

Main article: Camouflage

For a predator to locate a potential meal, it must first identify an organism as prey. Prey, however, have many adaptive characteristics which make such a task difficult. Crypsis is the ability of an animal, predator or prey, to make itself hard to see.[3]

Camouflage is one heavily utilised method, and involves the mimicking of the colour and patterns of the environment. This is achieved by external pigmentation patterns, and resting immobile and silent on an appropriate substrate, as do Egyptian Nightjar. Many marine animals use countershading or counterillumination to reduce their visibility from above and below. Some, such as the Mimic Octopus, can rapidly change their pattern and colour to provide a better defense. Some animals provide structures on their bodies for plant life to grow, camouflaging its host. Other animals masquerade as specific, inedible objects: the Bird Dropping Spider and the Phobaeticus Stick insects are notable examples.[4]

File:Polymorphism in Cepaea nemoralis.jpg
Polymorphism in the Grove snail is partly a response to predation by Song Thrushes

Visual Polymorphism, the existence of different forms within a single species, is a strategy used to reduce predation risk. Predators make use of search images to differentiate edible objects from all other objects; this causes frequency-dependent "apostatic selection" in which morphs with rarer colours and patterns are less likely to be killed. The striking polymorphism of the Grove snail may be partly caused by apostatic selection by predators including the Song Thrush.[5] There is evidence that polymorphic insect prey suffer less predation than single-morph species at a particular density, and can maintain higher population densities for a given rate of predation.[6][7]:11. This effect is more pronounced in tropical areas than temperate ones due to more intense and consistent interactions among different organisms.[8]

Avoiding predators in space and time

Animals alter the period in which they are awake in order to avoid predators.[9] Generally, animals are either diurnal (active during the day), nocturnal (active during the night), or crepuscular (active during twilight) depending on food availability and predator prevalence. For example, Dipodomys merriami kangaroo rats become crepuscular instead of nocturnal on full moon nights, when predators could see them more easily.[9]

Anolis sagrei lizards are more arboreal (living in trees) on islands where ground-living predators exist.[9]

Highly mobile creatures such as seabirds migrate to avoid predators during the period in which they are most vulnerable, the breeding season, for example moving to offshore islands to establish colonies far from the reach of land predators.

Avoiding attack

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Having been detected by a predator, many animals attempt to signal to a predator that they are not worth eating. Some animals make use of aposematic signals, for example bright warning colouration, or sounds and smells, so as to advertise that they are poisonous.[10] These patterns are often convergent, with red or yellow coupled with black being widely recognised as dangerous. Batesian mimicry is the imitation by a harmless species of the warning signals of a harmful species directed at a common predator, as can be seen in a hoverfly mimicking a wasp.[11]

File:Automeris ioPCCP20040706-5706AB.jpg
The eyespots of Automeris io

Some animals, particularly gazelles, are known to stot, which, among other things, may advertise their unprofitability to predators.[12] The moth Automeris io possesses eyespots hidden on its hindwings. When under threat, the moth suddenly reveals these spots, aiming to startle its predator. Fearful of their own consumption, predators often retreat when startled. Eyespots are also used by some animals to trick a predator into thinking it has been spotted (when in fact, it may not have been), and so is less likely to succeed in pursuit.[13]

Avoiding capture

Many animals have highly developed senses of sight, smell, and hearing so that they can detect danger and escape. By frequently scanning and monitoring their surroundings, especially when in the open, prey can avoid attack by hoping to see a predator before it reaches the 'critical distance' for an attack. This is a standard defense mechanism for animals in open grasslands and prairies. It is also common for arboreal animals to scan both the ground around them for terrestrial predators, and the sky for aerial predators. Smaller animals may not venture too far from cover in burrows or the undergrowth, where they can quickly hide when danger approaches. Flight is of huge importance in the avoidance of predators in those species that possess it.[14]

Predator Recognition

Learned Predator Recognition is influenced by pre-learning growth rate. That is the conclusion that was reached from the research done by Maud Ferrari, Grant Brown, Gary Bortolotti and Douglas Chivers on whether information learned about predators was influenced by prey growth trajectory before and after learning.[15]

Safety in numbers

Animals that are the frequent target of predation often make use of 'safety in numbers'. This results in a situation where any one herd member is unlikely to be preyed upon, and in high populations, predator satiation is likely to occur. Grazing mammals often feed in social groups, also known as herds. Working as a group, a stalking predator is likely to be detected earlier, and when it attacks, the herd scatters, causing difficulty for the predator and allowing most, if not all, of the prey animals to escape. Prey animals may use alarm signals to alert other herd members when a predator is sighted or sensed. Animals usually have a breeding season, where all the members of the species spawn at the same time, in order to maximise their young's chance of survival. This is particularly pronounced in insects such as Magicicada and mayflies, where millions of individuals emerge from pupation on the same day.[16]

Fighting off predators

File:MuskOxen.jpg
Muskoxen in defensive formation, horns ready, and highly alert

Many animals use horns, claws, and teeth to fight off predators. Some can inject venom and toxins, and skunks and bombardier beetles[17] spray noxious chemicals to deter attackers. Mobbing, the harassing of a predator by many prey animals, is common in birds, and is usually done to protect the young in social colonies. The Eastern Honeybee mobs invading hornets, vibrating their flight muscles in order to raise the temperature around the hornet scout to lethal levels, rather than allowing the scout to bring others to their beehive.[18]

Confusing the predator

File:Zebras Serengeti.JPG
A single zebra is hard to catch amongst a herd

Some animals have evolved dazzle camouflage, whereby instead of attempting to conceal themselves, they are patterned to cause motion dazzle, confusing a predator during an attack, and making it harder to select and track a target. An example is zebras, which stand out in the savannah when stationary, but when moving rapidly en masse, their stripes create a confusing, flickering mass in the eye of a predator such as a lion.[19]

Avoiding consumption

File:Fishing spider autotomy.jpg
This fishing spider has shed two of its legs to escape a predator

Having been captured, an animal must prevent the predator from killing and eating it. Mechanical defenses in those that possess them, such as armour and spines prevent access to softer edible parts. Distraction displays are used to direct the attention of the predator away from some vital area, such as the head, or a nest of chicks.[20] These can be visual, as in misdirecting eyespots, behavioural, such as a mother-bird feigning injury, or chemical, as seen in the production of ink clouds by squid and octopuses.

Some animals (e.g. possums) freeze in cover or play dead when seen. Often this is accompanied by foul smells, as if their corpse is in an advanced state of decomposition, one which many animals would avoid consuming.[21]

Some animals take a more drastic approach to defending themselves. Autotomy, the shedding of a non-vital bodypart, is utilised by some species to escape the grasp of a predator. Many lizards shed their tails when clasped, and arthropods will readily give up several legs if it allows their escape. When under threat, sea cucumbers rapidly eviscerate, ejecting part or all of their digestive tract. This is done to either anchor the cucumber into a rock fissure, or to eject toxins at the predator.[22] Horned lizards, when threatened, increase the pressure in their sinus cavities until the blood vessels in the corners of the eyes burst, squirting blood at the attacker.[23] Armoured crickets and many other insect species also use this method, known as autohaemorrhaging.[24] These animals are able to survive the loss of these tissues, and later regenerate them.

The soldier ant caste of the Malaysian species Camponotus saundersi undergoes a process known as autothysis to defend their ant colony. The soldier ants have two large glands that run the entire length of their body, and when stressed during battle, abdominal muscles contract, causing the glands to explode, killing the ant, but spraying poison in all directions.[25][26]

Plant adaptations

File:Aphid-sap.jpg
Aphid feeding on plant sap
Main article: Plant defense against herbivory

Many plant species have, over the course of their evolutionary history, developed physical and chemical[27] defense mechanisms to deter herbivores. Thorns, spines, and prickles are examples of physical mechanisms. Stinging nettles for instance are covered in hollow hairs that can inject irritant chemicals. Acacias and roses are among the many plants protected by spines and thorns. Prickles and spines are not effective against small insects such as aphids however, so some plants, such as some of the Solanaceae, supplement a bristly surface with sticky secretions that trap and kill small pests. Other plants such as holly have thick tough leaves covered with slippery way, making feeding difficult for small herbivores.[28]

Many Ericas have a sticky perianth, and some Moraeas have sticky peduncles that deter ants from plundering the flowers' nectar without contributing to their pollination. Many plants deter their enemies with repellent tastes or irritating or dangerous poisons in sap or in latex.[29] The herbivores in turn developed wide ranges of counter-adaptations.

See also

References

  1. John Alcock (1998). Animal Behavior: An Evolutionary Approach, 8th, Sinauer.
  2. Endler (1991) In Behavioural Ecology, 3rd ed. (Krebs & Davies), pp. 169–196.
  3. Forbes, Peter (2009). Dazzled and Deceived: Mimicry and Camouflage, 50-51, Yale.
  4. Cott, H.B. (1940) Adaptive Coloration in Animals. Methuen, London. (Stick insects: 334-335. Bird dropping spider, 330-332.)
  5. Cain, A.J. and Sheppard, P.M. (1954). Natural Selection in Cepaea. Genetics 39 (1): 89–116.
  6. Fullick & Greenwood (1979) Am. Nat. 113, 762-765.
  7. Edmunds, Malcolm. The Evolution of Cryptic Colour. in Insect defenses: adaptive mechanisms and strategies of prey and predators (1990) Eds. David L. Evans, Justin O. Schmidt, pg 11.
  8. Masaki Hoso, Michio Hori The American Naturalist, Vol. 172, No. 5 (Nov., 2008), pp. 726-732
  9. 9.0 9.1 9.2 Rosier and Langkilde, 2011
  10. Juan Carlos Santos, Luis A. Coloma, David C. Cannatella. Multiple, recurring origins of aposematism and diet specialization in poison frogs. National Academy of Sciences. URL accessed on 2008-12-22.
  11. Cott, H.B. (1940) Adaptive Coloration in Animals. Methuen, London. pages 396-413.
  12. Caro, T. M. (1986). The functions of stotting in Thomson's gazelles: Some tests of the predictions.. Animal Behaviour (34): 663–684.
  13. Stevens, Martin (2005). The role of eyespots as anti-predator mechanisms, principally demonstrated in the Lepidoptera. Biological Reviews 80 (4): 573–588.
  14. Rosier, Renee L; Langkilde, Tracy. Behavior Under Risk: How Animals Avoid Becoming Dinner. Nature. URL accessed on December 09, 2012.
  15. Ferrari, Maud C. O., Grant E. Brown • Gary R. Bortolotti • Douglas P. Chivers (November 2011). Prey behaviour across antipredator adaptation types: how does growth trajectory influence learning of predators?. Animal Cognition: 809–816.
  16. John Cooley & Dave Marshall. Periodical Cicada. University of Michigan. URL accessed on 2008-12-22.
  17. Piper, Ross (2007). Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals, Greenwood Press.
  18. Masato Ono, Takeshi Igarashi, Eishi Ohno, and Masami Sasaki (28 September 1995). Unusual thermal defence by a honeybee against mass attack by hornets. Nature 377 (377): 334–336.
  19. Martin Stevens, William TL Searle, Jenny E Seymour, Kate LA Marshall, Graeme D Ruxton. BMC Biology: Motion dazzle. Motion dazzle and camouflage as distinct anti-predator defenses. BMC Biology. URL accessed on January 5, 2012.
  20. Ruxton, Graeme D; Thomas N. Sherratt; Michael Patrick Speed. (2004) Avoiding attack: the evolutionary ecology of crypsis, warning signals and mimicry. Oxford University Press. ISBN 0-19-852859-0. p. 198
  21. Pasteur, G. (1982). "A classificatory review of mimicry systems". Annual Review of Ecology and Systematics 13: 169–199.
  22. Patrick Flammang, Jerome Ribesse, Michel Jangoux (2002-12-01). Biomechanics of adhesion in sea cucumber cuvierian tubules (echinodermata, holothuroidea). Integrative and Comparative Biology.
  23. Dr. Wendy Hodges. About Horned Lizards. DigiMorph. URL accessed on 2008-12-22.
  24. "See It to Believe It: Animals Vomit, Spurt Blood to Thwart Predators", Allison Bond, Discover Magazine blog, 28 July 2009, retrieved 17 March 2010
  25. Maschwitz, U. and E. Maschwitz, 1974. Platzende Arbeiterinnen: Eine neue Art der Feindabwehr bei sozialen Hautflüglern. Oecologia Berlin 14:289–294 (in German)
  26. C. Bordereau, A. Robert, V. Van Tuyen & A. Peppuy (1997). Suicidal defensive behavior by frontal gland dehiscence in Globitermes sulphureus Haviland soldiers (Isoptera). Insectes Sociaux 44 (3): 289–297.
  27. Biochemical defenses: secondary metabolites:. Plant Defense Systems & Medicinal Botany. URL accessed on 2007-05-21.
  28. Fernandes GW (1994). Plant mechanical defenses against insect herbivory. Revista Brasileira de Entomologia 38 (2): 421–433 [1].
  29. van Wyk, Ben-Erik; van Heerden, Fanie; van Oudtshoorn, Bosch (2002). Poisonous Plants of South Africa, Pretoria: Briza.

Further reading

  • Caro, T. 2005. Antipredator Defenses in Birds and Mammals. Chicago : University of Chicago Press. 591 pp. ISBN 0-226-09435-9 (hardcover version)
  • Cott, H.B. 1940. Adaptive Coloration in Animals. London: Methuen.
  • Edmunds, M. 1974. Defence in Animals: A Survey of Anti-Predator Defences. Harlow, Essex & NY: Longman ISBN 0-582-44132-3
  • Gabrielsen, G.W. & Smith, E.N.: Physiological responses associated with feigned death in the American Opossum. Acta Phys. Scand. 123: 393-398. 1985.
  • Gabrielsen, G.W. & Smith, E.N.: Physiological responses to disturbance in animals. In; Wildlife and Recreationists (R. Knight and K. Utzwiller. Island Press, Washington, D.C. pp. 137–153. 1995.
  • Rosier, R.L. & Langkilde, T.: Behavior Under Risk: How Animals Avoid Becoming Dinner. Nature Education Knowledge 2(11):8. 2011. Full text
  • Ruxton, G. D.; Speed, M. P.; Sherratt, T. N. (2004). Avoiding Attack. The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry. Oxford: Oxford University Press. ISBN 0-19-852860-4
  • Steen, J.B., Gabrielsen, G.W. & Kanwischer, J.W.: Physiological aspects of freezing behavior in Willow Ptarmigan hens. Acta Phys. Scand. 134: 299-304. 1988.


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