View source for Predation

{{short description|Biological interaction}}
{{Redirect2|Predator|Prey|other uses|Predator (disambiguation)|and|Prey (disambiguation)}}
{{good article}}
{{Use dmy dates|date=April 2024}}
{{Use British English|date=October 2025}}
[[File:Polar bear (Ursus maritimus) with its prey.jpg|thumb|upright=1.5|Solitary predator: a [[polar bear]] feeds on a [[bearded seal]] it has killed.]]
[[File:Ants eating cicada, jjron 22.11.2009.jpg|thumb|upright=1.5|[[Social predator]]s: [[meat ant]]s cooperate to feed on a [[cicada]] far larger than themselves.]] 

'''Predation''' is a [[biological interaction]] in which one organism, the '''predator''', kills and eats another organism, its '''prey'''. It is one of a family of common [[List of feeding behaviours|feeding behaviours]] that includes [[parasitism]] and [[micropredation]] (which usually do not kill the [[Host (biology)|host]]) and [[parasitoid]]ism (which always does, eventually). It is distinct from [[Scavenger|scavenging]] on dead prey, though many predators also scavenge; it overlaps with [[Herbivore|herbivory]], as [[Seed predation|seed predators]] and destructive [[frugivore]]s are predators.

Predation behaviour varies significantly depending on the organism. Many predators, especially [[carnivore]]s, have evolved distinct [[hunting strategy|hunting strategies]]. [[Pursuit predation]] involves the active search for and pursuit of prey, whilst [[ambush predation|ambush predators]] instead wait for prey to present an opportunity for capture, and often use stealth or [[aggressive mimicry]]. Other predators are [[opportunism|opportunistic]] or [[omnivore|omnivorous]] and only practice predation occasionally.

Most [[Carnivore#Obligate carnivores|obligate carnivores]] are specialized for hunting. They may have acute senses such as [[eye|vision]], [[hearing]], or [[olfaction|smell]] for [[prey detection]]. Many predatory animals have sharp [[claw]]s or [[jaw]]s to grip, kill, and cut up their prey. [[Physical strength]] is usually necessary for large carnivores such as [[big cat]]s to kill larger prey. Other adaptations include stealth, [[endurance]], [[intelligence]], [[social behaviour]], and aggressive mimicry that improve hunting efficiency.

Predation has a powerful [[selection pressure|selective effect]] on prey, and the prey develops [[anti-predator adaptation]]s such as [[warning coloration]]<!--yes, in BE as well as AE-->, [[alarm call]]s and other [[signalling theory|signals]], [[camouflage]], [[mimicry]] of well-defended species, and defensive spines and chemicals. Sometimes predator and prey find themselves in an [[evolutionary arms race]], a cycle of adaptations and counter-adaptations. Predation has been a major driver of [[evolution]] since at least the [[Cambrian]] period.

==Definition==
[[File:IndianSpiderWasp.JPG|thumb|upright|left|[[Pompilidae|Spider wasps]] paralyse and eventually kill their hosts, but are considered [[parasitoid]]s, not predators.]]

At the most basic level, predators kill and eat other organisms. However, the concept of predation is broad, defined differently in different contexts, and includes a wide variety of feeding methods; moreover, some relationships that result in the prey's death are not necessarily called predation. A [[parasitoid]], such as an [[ichneumon wasp]], lays its eggs in or on its host; the eggs hatch into larvae, which eat the host, and it inevitably dies. Zoologists generally call this a form of [[parasitism]], though conventionally parasites are thought not to kill their hosts. A predator can be defined to differ from a parasitoid in that it has many prey, captured over its lifetime, where a parasitoid's larva has just one, or at least has its food supply provisioned for it on just one occasion.<ref>{{cite book |last1=Gurr |first1=Geoff M. |last2=Wratten |first2=Stephen D. |last3=Snyder |first3=William E. |title=Biodiversity and Insect Pests: Key Issues for Sustainable Management |url=https://books.google.com/books?id=-ng4hIDQ_k8C&pg=PT105 |year=2012 |publisher=John Wiley & Sons |isbn=978-1-118-23185-2 |page=105}}</ref><ref name=LaffertyKuris2002/>

[[File:Predation's Boundaries.svg|thumb|upright=1.5|Relation of predation to other feeding strategies]]

There are other difficult and borderline cases. [[Micropredator]]s are small animals that, like predators, feed entirely on other organisms; they include [[flea]]s and [[mosquito]]es that consume blood from living animals, and [[aphid]]s that consume sap from living plants. However, since they typically do not kill their hosts, they are now often thought of as parasites.<ref name=PoulinRandhawa2015>{{cite journal |last1=Poulin |first1=Robert |author1-link=Robert Poulin (zoologist) |last2=Randhawa |first2=Haseeb S. |title=Evolution of parasitism along convergent lines: from ecology to genomics |journal=Parasitology |date=February 2015 |volume=142 |issue=Suppl 1 |pages=S6–S15 |doi=10.1017/S0031182013001674 |pmc=4413784 |pmid=24229807}}</ref><ref name=Rollinson>{{cite book |doi=10.1016/B978-0-12-385897-9.00001-X |chapter=The Many Roads to Parasitism |title=Advances in Parasitology Volume 74 |date=2011 |last1=Poulin |first1=Robert |volume=74 |pages=1–40 |pmid=21295676 |isbn=978-0-12-385897-9 }}</ref> Animals that [[Grazing|graze]] on [[phytoplankton]] or mats of microbes are predators, as they consume and kill their food organisms, while herbivores that browse leaves are not, as their food plants usually survive the assault.<ref name="Bengtson2002"/> When animals eat seeds (''[[seed predation]]'' or ''granivory'') or eggs (''[[egg predation]]''), they are consuming entire living organisms, which by definition makes them predators.<ref name=Janzen>{{cite journal |last1=Janzen |first1=D. H. |year=1971 |title=Seed Predation by Animals |journal=Annual Review of Ecology and Systematics |volume=2 |issue=1 |pages=465–492 |doi=10.1146/annurev.es.02.110171.002341|bibcode=1971AnRES...2..465J }}</ref><ref name="NilssonBjörkman1985">{{cite journal |last1=Nilsson |first1=Sven G. |last2=Björkman |first2=Christer |last3=Forslund |first3=Pär |last4=Höglund |first4=Jacob |title=Egg predation in forest bird communities on islands and mainland |journal=Oecologia |volume=66 |issue=4 |year=1985 |pages=511–515 |doi=10.1007/BF00379342|pmid=28310791 |bibcode=1985Oecol..66..511N |s2cid=2145031 }}</ref><ref name=HulmeBenkman2002/>

[[Scavenger]]s, organisms that only eat organisms found already dead, are not predators, but many predators such as the [[jackal]] and the [[hyena]] scavenge when the opportunity arises.<ref>{{cite journal |last1=Kane |first1=Adam |last2=Healy |first2=Kevin |last3=Guillerme |first3=Thomas |last4=Ruxton |first4=Graeme D. |last5=Jackson |first5=Andrew L. |date=2017 |title=A recipe for scavenging in vertebrates – the natural history of a behaviour |journal=Ecography |volume=40 |issue=2 |pages=324–334 |doi=10.1111/ecog.02817 |bibcode=2017Ecogr..40..324K |hdl=10468/3213 |s2cid=56280901 |hdl-access=free }}</ref><ref>{{cite book |last=Kruuk |first=Hans |title=The Spotted Hyena: A Study of Predation and Social Behaviour |publisher=University of California Press |year=1972 |isbn=978-0-226-45508-2|pages=107–108}}</ref><ref name="Bengtson2002"/> Among invertebrates, [[social wasp]]s such as [[yellowjacket]]s are both hunters and scavengers of other insects.<ref name="Schmidt2009">{{cite book |doi=10.1016/B978-0-12-374144-8.00275-7 |chapter=Wasps |title=Encyclopedia of Insects |date=2009 |last1=Schmidt |first1=Justin O. |pages=1049–1052 |isbn=978-0-12-374144-8 }}</ref>

==Taxonomic range==
{{further|Carnivorous plant|Nematophagous fungus|Seed predation|Egg predation}}

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|caption1 = [[Carnivorous plant]]: [[sundew]] engulfing an insect
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|caption2 = [[Seed predation]]: [[African pygmy mouse|mouse]] eating seeds
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While examples of predators among mammals and birds are well known,<ref name=Stevens2010/> predators can be found in a broad range of taxa including arthropods. They are common among insects, including mantids, [[Dragonfly|dragonflies]], [[Neuroptera|lacewings]] and [[Mecoptera|scorpionflies]]. In some species such as the [[alderfly]], only the larvae are predatory (the adults do not eat). Spiders are predatory, as well as other terrestrial invertebrates such as [[scorpion]]s; [[centipede]]s; some [[mite]]s, [[snail]]s and [[slug]]s; [[nematode]]s; and [[Planarian|planarian worms]].<ref>{{cite web |title=Predators, parasites and parasitoids |url=https://australianmuseum.net.au/predators-parasites-and-parasitoids |website=Australian Museum |access-date=19 September 2018 |language=en}}</ref> In marine environments, most [[cnidaria]]ns (e.g., [[jellyfish]], [[hydroid (zoology)|hydroids]]), [[ctenophora]] (comb jellies), [[echinoderm]]s (e.g., [[sea stars]], [[sea urchin]]s, [[sand dollar]]s, and [[sea cucumber]]s) and [[flatworm]]s are predatory.<ref>{{cite encyclopedia |last1=Watanabe|first1=James M. |title=Invertebrates, overview |editor-last1=Denny |editor-first1=Mark W. |editor-last2=Gaines |editor-first2=Steven Dean |encyclopedia=Encyclopedia of tidepools and rocky shores |date=2007 |publisher=[[University of California Press]] |isbn=978-0-520-25118-2}}</ref> Among [[crustacean]]s, [[lobster]]s, [[crab]]s, [[shrimp]]s and [[barnacle]]s are predators,<ref>{{cite book |last1=Phelan |first1=Jay |title=What Is life?: a guide to biology |date=2009 |publisher=W.H. Freeman & Co |isbn=978-1-4292-2318-8 |page=432 |edition=Student}}</ref> and in turn crustaceans are preyed on by nearly all [[cephalopod]]s (including [[octopus]]es, [[squid]] and [[cuttlefish]]).<ref>{{cite journal |last1=Villanueva |first1=Roger |last2=Perricone |first2=Valentina |last3=Fiorito |first3=Graziano |title=Cephalopods as Predators: A Short Journey among Behavioral Flexibilities, Adaptions, and Feeding Habits |journal=Frontiers in Physiology |date=17 August 2017 |volume=8 |page=598 |doi=10.3389/fphys.2017.00598|pmid=28861006 |pmc=5563153 |doi-access=free }}</ref>

[[File:Инфузория туфелька поедает бактерии!.gif|thumb|''[[Paramecium]]'', a predatory [[ciliate]], feeding on [[bacteria]]]]

Seed predation is restricted to mammals, birds, and insects but is found in almost all terrestrial ecosystems.<ref name=HulmeBenkman2002>{{cite book |last1=Hulme |first1=P. E. |last2=Benkman |first2=C. W. |year=2002 |chapter =Granivory |pages=132–154 |title=Plant animal Interactions: An Evolutionary Approach |editor1=C. M. Herrera |editor2=O. Pellmyr |publisher=Blackwell |isbn=978-0-632-05267-7}}</ref><ref name=Janzen/> Egg predation includes both specialist egg predators such as some [[colubrid]] [[snake]]s and generalists such as foxes and badgers that opportunistically take eggs when they find them.<ref name="HanssenErikstad2012">{{cite journal |last1=Hanssen |first1=Sveinn Are |last2=Erikstad |first2=Kjell Einar |title=The long-term consequences of egg predation |journal=Behavioral Ecology |volume=24 |issue=2 |date=2012 |doi=10.1093/beheco/ars198 |pages=564–569|doi-access=free }}</ref><ref name="Pike Clark Manica Tseng p. ">{{cite journal |last1=Pike |first1=David A. |last2=Clark |first2=Rulon W. |last3=Manica |first3=Andrea |last4=Tseng |first4=Hui-Yun |last5=Hsu |first5=Jung-Ya |last6=Huang |first6=Wen-San |title=Surf and turf: predation by egg-eating snakes has led to the evolution of parental care in a terrestrial lizard |journal=Scientific Reports |volume=6 |issue=1 |article-number=22207 |date=26 February 2016 |doi=10.1038/srep22207|pmid=26915464 |pmc=4768160 |bibcode=2016NatSR...622207P }}</ref><ref>{{cite book |doi=10.13140/RG.2.1.1014.6960 |date=2016 |last1=Ainsworth |first1=Gillian B. |last2=Calladine |first2=John |last3=Martay |first3=Blaise |last4=Park |first4=Kirsty |last5=Redpath |first5=Steve |last6=Wernham |first6=Chris |last7=Wilson |first7=Mark |last8=Young |first8=Juliette |title=Understanding Predation: A review bringing together natural science and local knowledge of recent wild bird population changes and their drivers in Scotland |publisher=Scotland's Moorland Forum}}</ref>

Some plants, like the [[pitcher plant]], the [[Venus fly trap]] and the [[sundew]], are [[carnivorous plant|carnivorous and consume insects]].<ref name=Stevens2010/> Methods of predation by plants varies greatly but often involves a food trap, mechanical stimulation, and electrical impulses to eventually catch and consume its prey.<ref>{{cite journal |last1=Hedrich |first1=Rainer |last2=Fukushima |first2=Kenji |date=20 May 2021 |title=On the Origin of Carnivory: Molecular Physiology and Evolution of Plants on an Animal Diet |journal=Annual Review of Plant Biology |volume=72 |issue=1 |at=annurev–arplant–080620-010429 |doi=10.1146/annurev-arplant-080620-010429 |pmid=33434053 |s2cid=231595236 |issn=1543-5008|doi-access=free |bibcode=2021AnRPB..72..133H }}</ref> Some [[nematophagous fungus|carnivorous fungi]] catch [[nematode]]s using either active traps in the form of constricting rings, or passive traps with adhesive structures.<ref>{{cite journal |author=Pramer, D. |year=1964 |title=Nematode-trapping fungi |journal=Science |volume=144 |issue=3617 |pages=382–388 |jstor=1713426|doi=10.1126/science.144.3617.382 |pmid=14169325 |bibcode=1964Sci...144..382P }}</ref>

Many species of [[protozoa]] ([[eukaryote]]s) and [[bacteria]] ([[prokaryote]]s) prey on other microorganisms; the feeding mode is evidently ancient, and evolved many times in both groups.<ref name=VelicerMendes-Soares2007/><ref name=Stevens2010/><ref name="JurkevitchDavidov2006">{{cite book |last1=Jurkevitch |first1=Edouard |last2=Davidov |first2=Yaacov |title=Predatory Prokaryotes |url=https://archive.org/details/predatoryprokary00jurk |url-access=limited |chapter=Phylogenetic Diversity and Evolution of Predatory Prokaryotes |publisher=Springer |date=2006 |isbn=978-3-540-38577-6 |doi=10.1007/7171_052 |pages=[https://archive.org/details/predatoryprokary00jurk/page/n17 11]–56}}</ref> Among freshwater and marine [[zooplankton]], whether single-celled or multi-cellular, predatory grazing on [[phytoplankton]] and smaller zooplankton is common, and found in many species of [[nanoflagellate]]s, [[dinoflagellate]]s, [[ciliate]]s, [[rotifer]]s, a diverse range of [[meroplankton]] animal larvae, and two groups of crustaceans, namely [[copepod]]s and [[cladocera]]ns.<ref name="HansenBjørnsenHansen1997">{{cite journal |last1=Hansen |first1=Per Juel |last2=Bjørnsen |first2=Peter Koefoed |last3=Hansen |first3=Benni Winding |title=Zooplankton grazing and growth: Scaling within the 2-2,-μm body size range |journal=Limnology and Oceanography |volume=42 |issue=4 |year=1997 |doi=10.4319/lo.1997.42.4.0687 |pages=687–704 |bibcode=1997LimOc..42..687H |doi-access=free }} summarizes findings from many authors.</ref>

==Foraging==
{{See also|Foraging}}

[[File:Foraging Sequence.svg|thumb|upright=1.8|A basic [[foraging]] cycle for a predator, with some variations indicated<ref name=Kramer2001/>]]

To feed, a predator must search for, pursue and kill its prey. These actions form a [[foraging]] cycle.<ref name=Griffiths>{{cite journal |last1=Griffiths |first1=David |title=Foraging costs and relative prey size |journal=The American Naturalist |date=November 1980 |volume=116 |issue=5 |pages=743–752|jstor=2460632|doi=10.1086/283666 |bibcode=1980ANat..116..743G |s2cid=85094710 }}</ref><ref name="WetzelLikens2000">{{cite book |doi=10.1007/978-1-4757-3250-4_17 |chapter=Predator-Prey Interactions |title=Limnological Analyses |date=2000 |last1=Wetzel |first1=Robert G. |last2=Likens |first2=Gene E. |pages=257–262 |isbn=978-1-4419-3186-3 }}</ref> The predator must decide where to look for prey based on its geographical distribution; and once it has located prey, it must assess whether to pursue it or to wait for a better choice. If it chooses pursuit, its physical capabilities determine the mode of pursuit (e.g., ambush or chase).<ref name=Pianka>{{cite book |last1=Pianka |first1=Eric R. |title=Evolutionary ecology |date=2011 |publisher=Eric R. Pianka |pages=78–83 |edition=7th (eBook)}}</ref><ref>{{cite book |chapter=The economics of consumer choice |last1=MacArthur |first1=Robert H. |title=Geographical ecology: patterns in the distribution of species |date=1984 |publisher=Princeton University Press |isbn=978-0-691-02382-3 |pages=59–76}}</ref> Having captured the prey, it may also need to expend energy ''handling'' it (e.g., killing it, removing any shell or spines, and ingesting it).<ref name=Kramer2001>{{cite book |doi=10.1093/oso/9780195131543.003.0024 |chapter=Foraging Behavior |title=Evolutionary Ecology |date=2001 |last1=Kramer |first1=Donald L. |isbn=978-0-19-513154-3 }}</ref><ref name=Griffiths/>

===Search===

Predators have a choice of search modes ranging from ''sit-and-wait'' to ''active'' or ''widely foraging''.<ref name=Bell4>{{harvnb|Bell|2012|pages=4–5}}</ref><ref name=Kramer2001/><ref>{{cite book |last1=Eastman |first1=Lucas B. |last2=Thiel |first2=Martin |contribution=Foraging behavior of crustacean predators and scavengers |editor-last1=Thiel |editor-first1=Martin |editor-last2=Watling |editor-first2=Les |title=Lifestyles and feeding biology |date=2015 |publisher=Oxford University Press |isbn=978-0-19-979706-6 |pages=535–556}}</ref><ref>{{cite journal |last1=Perry |first1=Gad |title=The Evolution of Search Modes: Ecological versus Phylogenetic Perspectives |journal=The American Naturalist |date=January 1999 |volume=153 |issue=1 |pages=98–109 |doi=10.1086/303145|pmid=29578765 |bibcode=1999ANat..153...98P |s2cid=4334462 }}</ref> The sit-and-wait method is most suitable if the prey are dense and mobile, and the predator has low energy requirements.<ref name=Bell4/> Wide foraging expends more energy, and is used when prey is sedentary or sparsely distributed.<ref name=Pianka/><ref name=Bell4/> There is a continuum of search modes with intervals between periods of movement ranging from seconds to months. Sharks, [[Molidae|sunfish]], [[Insectivorous]] birds and [[shrew]]s are almost always moving while web-building spiders, aquatic invertebrates, praying mantises and [[kestrel]]s rarely move. In between, [[plover]]s and other [[shorebirds]], freshwater fish including [[crappie]]s, and the larvae of [[Coccinellidae|coccinellid beetles (ladybirds)]], alternate between actively searching and scanning the environment.<ref name=Bell4/>

[[File:Thalassarche melanophrys in flight 2 - SE Tasmania.jpg|thumb|left|The [[black-browed albatross]] regularly flies hundreds of kilometres across the nearly empty ocean to find patches of food.]]

Prey distributions are often clumped, and predators respond by looking for ''patches'' where prey is dense and then searching within patches.<ref name=Kramer2001/> Where food is found in patches, such as rare shoals of fish in a nearly empty ocean, the search stage requires the predator to travel for a substantial time, and to expend a significant amount of energy, to locate each food patch.<ref name=Bell2012/> For example, the [[black-browed albatross]] regularly makes foraging flights to a range of around {{convert|700|km|mi|abbr=off|-1}}, up to a maximum foraging range of {{convert|3000|km|mi|abbr=off|-1}} for breeding birds gathering food for their young.{{efn|A range of 3000 kilometres means a flight distance of at least 6000 kilometres out and back.}}<ref name="Gremillet2000">{{cite journal |last1=Gremillet |first1=D. |last2=Wilson |first2=R. P. |last3=Wanless |first3=S. |last4=Chater |first4=T. |title=Black-browed albatrosses, international fisheries and the Patagonian Shelf |journal=Marine Ecology Progress Series |date=2000 |volume=195 |pages=69–280|doi=10.3354/meps195269 |bibcode=2000MEPS..195..269G |doi-access=free }}</ref> With static prey, some predators can learn suitable patch locations and return to them at intervals to feed.<ref name=Bell2012>{{harvnb|Bell|2012|pages=69–188}}</ref> The [[optimal foraging theory|optimal foraging]] strategy for search has been modelled using the [[marginal value theorem]].<ref name="Charnov1976">{{cite journal |last=Charnov |first=Eric L. |title=Optimal foraging, the marginal value theorem |journal=Theoretical Population Biology |volume=9 |issue=2 |year=1976 |doi=10.1016/0040-5809(76)90040-x |pmid=1273796 |pages=129–136 |bibcode=1976TPBio...9..129C }}</ref>

Search patterns often appear random. One such is the [[Lévy flight|Lévy walk]], that tends to involve clusters of short steps with occasional long steps. It is a [[Lévy flight foraging hypothesis|good fit to the behaviour]] of a wide variety of organisms including bacteria, honeybees, sharks and human hunter-gatherers.<ref>{{cite journal |last1=Reynolds |first1=Andy |title=Liberating Lévy walk research from the shackles of optimal foraging |journal=Physics of Life Reviews |date=September 2015 |volume=14 |pages=59–83 |doi=10.1016/j.plrev.2015.03.002 |pmid=25835600 |bibcode=2015PhLRv..14...59R }}</ref><ref>{{cite journal |last1=Buchanan |first1=Mark |title=Ecological modelling: The mathematical mirror to animal nature |journal=Nature |date=5 June 2008 |volume=453 |issue=7196 |pages=714–716 |doi=10.1038/453714a |pmid=18528368 |doi-access=free }}</ref>

===Assessment===

[[File:Ladybug aphids.JPG|thumb|[[Seven-spot ladybird]]s select plants of good quality for their [[aphid]] prey.]]

Having found prey, a predator must decide whether to pursue it or keep searching. The decision depends on the costs and benefits involved. A bird foraging for insects spends a lot of time searching but capturing and eating them is quick and easy, so the efficient strategy for the bird is to eat every palatable insect it finds. By contrast, a predator such as a lion or falcon finds its prey easily but capturing it requires a lot of effort. In that case, the predator is more selective.<ref name=Pianka/>

One of the factors to consider is size. Prey that is too small may not be worth the trouble for the amount of energy it provides. Too large, and it may be too difficult to capture. For example, a mantid captures prey with its forelegs and they are optimized for grabbing prey of a certain size. Mantids are reluctant to attack prey that is far from that size. There is a positive correlation between the size of a predator and its prey.<ref name=Pianka/>

A predator may assess a patch and decide whether to spend time searching for prey in it.<ref name=Kramer2001/> This may involve some knowledge of the preferences of the prey; for example, [[ladybird]]s can choose a patch of vegetation suitable for their [[aphid]] prey.<ref name="WilliamsFlaxman2012">{{cite journal |last1=Williams |first1=Amanda C. |last2=Flaxman |first2=Samuel M. |title=Can predators assess the quality of their prey's resource? |journal=Animal Behaviour |volume=83 |issue=4 |year=2012 |doi=10.1016/j.anbehav.2012.01.008 |pages=883–890 |s2cid=53172079}}</ref>

===Capture===

To capture prey, predators have a spectrum of pursuit modes that range from overt chase (''[[pursuit predation]]'') to a sudden strike on nearby prey (''[[ambush predator|ambush predation]]'').<ref name=Kramer2001/><ref>{{cite journal |last1=Scharf |first1=Inon |last2=Nulman |first2=Einat |last3=Ovadia |first3=Ofer |last4=Bouskila |first4=Amos |date=September 2006 |title=Efficiency evaluation of two competing foraging modes under different conditions |journal=The American Naturalist |volume=168 |issue=3 |pages=350–357 |doi=10.1086/506921 |pmid=16947110|bibcode=2006ANat..168..350S |s2cid=13809116 }}</ref><ref name=Stevens2010>{{cite journal |author=Stevens, Alison N. P. |date=2010 |title=Predation, Herbivory, and Parasitism |journal=Nature Education Knowledge |volume=3 |issue=10 |page=36 |url=https://www.nature.com/scitable/knowledge/library/predation-herbivory-and-parasitism-13261134}}</ref> Another strategy in between ambush and pursuit is ''ballistic interception'', where a predator observes and predicts a prey's motion and then launches its attack accordingly.<ref name="MooreBiewener2015"/>

====Ambush====
{{main|Ambush predation}}

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| caption1 = [[Western green lizard]] [[ambush predator|ambushes]] its [[grasshopper]] prey.
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Ambush or sit-and-wait predators are carnivorous animals that capture prey by stealth or surprise. In animals, ambush predation is characterized by the predator's scanning the environment from a concealed position until a prey is spotted, and then rapidly executing a fixed surprise attack.<ref name=deVries>{{cite journal |author=deVries, M. S. |author2=Murphy, E. A. K. |author3=Patek S. N. |title=Strike mechanics of an ambush predator: the spearing mantis shrimp |year=2012 |journal=Journal of Experimental Biology |volume=215 |issue=Pt 24 |pages=4374–4384 |doi=10.1242/jeb.075317 |pmid=23175528 |doi-access=free |bibcode=2012JExpB.215.4374D }}</ref><ref name="MooreBiewener2015"/> Vertebrate ambush predators include frogs, fish such as the [[angel shark]], the [[northern pike]] and the [[eastern frogfish]].<ref name="MooreBiewener2015"/><ref name="WhosWho">{{cite web |url=http://www.hww.ca/hww2.asp?pid=1&id=87&cid=8 |archive-url=https://web.archive.org/web/20070518235841/http://www.hww.ca/hww2.asp?pid=1&id=87&cid=8 |archive-date=18 May 2007 |title=Cougar |website=Hinterland Who's Who |access-date=22 May 2007 |publisher=[[Canadian Wildlife Service]] and [[Canadian Wildlife Federation]]}}</ref><ref>{{cite web |title=Pikes (Esocidae) |url=https://www.in.gov/dnr/fishwild/files/fw-pikes.pdf |publisher=Indiana Division of Fish and Wildlife |access-date=3 September 2018}}</ref><ref name=fishes>{{cite web |last1=Bray |first1=Dianne |title=Eastern Frogfish, Batrachomoeus dubius |url=https://fishesofaustralia.net.au/home/species/2835 |website=Fishes of Australia |access-date=14 September 2014 |url-status=live |archive-url=https://web.archive.org/web/20140914181407/http://www.fishesofaustralia.net.au/home/species/2835 |archive-date=14 September 2014 }}</ref> Among the many invertebrate ambush predators are [[trapdoor spider]]s and [[Thomisus spectabilis|Australian Crab spiders]] on land and [[mantis shrimp]]s in the sea.<ref name=deVries/><ref>{{cite web |url=https://www.bbc.co.uk/nature/life/Liphistius |archive-url=https://web.archive.org/web/20101203042945/http://www.bbc.co.uk/nature/life/Liphistius |archive-date=3 December 2010 |title=Trapdoor spiders |publisher=BBC |access-date=12 December 2014}}</ref><ref>{{cite web |url=http://www.desertmuseum.org/books/nhsd_trapdoor_spider.php|access-date=12 December 2014 |title=Trapdoor spider |publisher=[[Arizona-Sonora Desert Museum]] |year=2014}}</ref> Ambush predators often construct a burrow in which to hide, improving concealment at the cost of reducing their field of vision. Some ambush predators also use lures to attract prey within striking range.<ref name="MooreBiewener2015"/> The capturing movement has to be rapid to trap the prey, given that the attack is not modifiable once launched.<ref name="MooreBiewener2015"/>

{{anchor|Ballistic}}

====Ballistic interception====

{{multiple image
| align    = right
| image1   = Chameleon gab fbi.png
| width1   = 220
| caption1 = Tongue, adapted for prey capture
| image2   = Chameleon-Stage 03.jpg
| width2   = 185
| caption2 = Capturing a fly at a distance
| footer   = The [[chameleon]] attacks prey by shooting out its sticky tongue.
}}

Ballistic interception is the strategy where a predator observes the movement of a prey, predicts its motion, works out an interception path, and then attacks the prey on that path. This differs from ambush predation in that the predator adjusts its attack according to how the prey is moving.<ref name="MooreBiewener2015"/> Ballistic interception involves a brief period for planning, giving the prey an opportunity to escape. Some frogs wait until snakes have begun their strike before jumping, reducing the time available to the snake to recalibrate its attack, and maximising the angular adjustment that the snake would need to make to intercept the frog in real time.<ref name="MooreBiewener2015"/> Ballistic predators include insects such as dragonflies, and vertebrates such as [[archerfish]] (attacking with a jet of water), [[chameleon]]s (attacking with their tongues), and some [[Colubridae|colubrid snakes]].<ref name="MooreBiewener2015">{{cite journal |last1=Moore |first1=Talia Y. |last2=Biewener |first2=Andrew A. |title=Outrun or Outmaneuver: Predator–Prey Interactions as a Model System for Integrating Biomechanical Studies in a Broader Ecological and Evolutionary Context |journal=Integrative and Comparative Biology |volume=55 |issue=6 |pages=1188–97 |date=2015 |doi=10.1093/icb/icv074 |pmid=26117833 |doi-access=free}}</ref>

====Pursuit====
{{main|Pursuit predation}}

{{multiple image
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| image1 = Whales Bubble Net Feeding-edit1.jpg
| width1 = 200
| alt1 = 
| caption1 = [[Humpback whale]]s are lunge feeders, filtering thousands of [[krill]] from seawater and swallowing them alive.
| image2 = Gomphus vulgatissimus with a prey 002.jpg
| width2 = 188
| alt2 = 
| caption2 = [[Dragonfly|Dragonflies]], like this [[Gomphus vulgatissimus|common clubtail]] with captured prey, are invertebrate [[Pursuit predation|pursuit predators]].
| footer = 
}}

In pursuit predation, predators chase fleeing prey. If the prey flees in a straight line, capture depends only on the predator's being faster than the prey.<ref name="MooreBiewener2015"/> If the prey manoeuvres by turning as it flees, the predator must react in real time to calculate and follow a new intercept path, such as by [[parallel navigation]], as it closes on the prey.<ref name="MooreBiewener2015"/> Many pursuit predators use camouflage to approach the prey as close as possible unobserved (''stalking'') before starting the pursuit.<ref name="MooreBiewener2015"/> Pursuit predators include terrestrial mammals such as humans, African wild dogs, spotted hyenas and wolves; marine predators such as dolphins, orcas and many predatory fishes, such as tuna;<ref>{{cite journal |last1=Gazda |first1=S. K. |last2=Connor |first2=R. C. |last3=Edgar |first3=R. K. |last4=Cox |first4=F. |year=2005 |title=A division of labour with role specialization in group-hunting bottlenose dolphins (Tursiops truncatus) off Cedar Key, Florida |journal=Proceedings of the Royal Society |volume=272 |issue=1559 |pages=135–140 |doi=10.1098/rspb.2004.2937 |pmid=15695203 |pmc=1634948}}</ref><ref>{{cite book |last=Tyus |first=Harold M. |title=Ecology and Conservation of Fishes |url=https://books.google.com/books?id=K2fNBQAAQBAJ&pg=PA233 |year=2011 |publisher=CRC Press |isbn=978-1-4398-9759-1 |page=233}}</ref> predatory birds (raptors) such as falcons; and insects such as [[dragonfly|dragonflies]].<ref name=Combes>{{cite journal |last1=Combes |first1=S. A. |last2=Salcedo |first2=M. K. |last3=Pandit |first3=M. M. |last4=Iwasaki |first4=J. M. |year=2013 |title=Capture Success and Efficiency of Dragonflies Pursuing Different Types of Prey |journal=Integrative and Comparative Biology |volume= 53 |issue=5 |pages=787–798 |doi=10.1093/icb/ict072 |pmid=23784698 |doi-access=free }}</ref>

An extreme form of pursuit is [[Persistence hunting|endurance or persistence hunting]], in which the predator tires out the prey by following it over a long distance, sometimes for hours at a time. The method is used by human [[hunter-gatherer]]s and by [[Canidae|canids]] such as [[African wild dog]]s and domestic hounds. The African wild dog is an extreme persistence predator, tiring out individual prey by following them for many miles at relatively low speed.<ref name="HubelMyattJordanDewhirst2016">{{cite journal |last1=Hubel |first1=Tatjana Y. |last2=Myatt |first2=Julia P. |last3=Jordan |first3=Neil R. |last4=Dewhirst |first4=Oliver P. |last5=McNutt |first5=J. Weldon |last6=Wilson |first6=Alan M. |title=Energy cost and return for hunting in African wild dogs |journal=Nature Communications |volume=7 | date=29 March 2016 |doi=10.1038/ncomms11034 |pmid=27023457 |pmc=4820543 | article-number=11034 |quote=Cursorial hunting strategies range from one extreme of transient acceleration, power and speed to the other extreme of persistence and endurance with prey being fatigued to facilitate capture.Dogs and humans are considered to rely on endurance rather than outright speed and manoeuvrability for success when hunting cursorially.}}</ref>

A specialised form of pursuit predation is the [[lunge feed]]ing of [[baleen whale]]s. These very large marine predators feed on [[plankton]], especially [[krill]], diving and actively swimming into concentrations of plankton, and then taking a huge gulp of water and [[Filter feeder|filtering]] it through their feathery [[baleen]] plates.<ref>{{cite journal |last1=Goldbogen |first1=J. A. |last2=Calambokidis |first2=J. |last3=Shadwick |first3=R. E. |last4=Oleson |first4=E. M. |last5=McDonald |first5=M. A. |last6=Hildebrand |first6=J. A. |year=2006 |title=Kinematics of foraging dives and lunge-feeding in fin whales |journal=Journal of Experimental Biology |volume=209 |issue=7 |pages=1231–1244 |doi=10.1242/jeb.02135 |pmid=16547295 | s2cid=17923052 |doi-access=free |bibcode=2006JExpB.209.1231G }}</ref><ref>{{cite journal |last1=Sanders |first1=Jon G. |last2=Beichman |first2=Annabel C. |last3=Roman |first3=Joe |last4=Scott |first4=Jarrod J. |last5=Emerson |first5=David |last6=McCarthy |first6=James J. |last7=Girguis |first7=Peter R. |author7-link=Peter Girguis |year=2015 |title=Baleen whales host a unique gut microbiome with similarities to both carnivores and herbivores |journal=Nature Communications |volume=6 |article-number=8285 |doi=10.1038/ncomms9285 |pmid=26393325 |pmc=4595633 |bibcode=2015NatCo...6.8285S}}</ref>

Pursuit predators may be [[Social predator|social]], like the lion and wolf that hunt in groups, or solitary.<ref name=LaffertyKuris2002/><!--Ambush predators are often solitary.-->

===Handling===

{{multiple image
| align             = right
| image1            = Bluecat5A.jpg
| width1            = 225
| caption1          = [[Catfish]] have sharp [[fish fin|dorsal and pectoral spines]] which are held erect to discourage predators such as [[heron]]s which swallow prey whole.
| image2            = Osprey eating a fish.jpg
| width2            = 165
| caption2          = [[Osprey]] tears its fish prey apart, avoiding dangers such as sharp spines.
}}

Once the predator has captured the prey, it has to handle it: very carefully if the prey is dangerous to eat, such as if it possesses sharp or poisonous spines, as in many prey fish. Some [[catfish]] such as the [[Ictaluridae]] have [[fish fin|spines on the back (dorsal) and belly (pectoral)]] which lock in the erect position; as the catfish thrashes about when captured, these could pierce the predator's mouth, possibly fatally. Some fish-eating birds like the [[osprey]] avoid the danger of spines by tearing up their prey before eating it.<ref name="Forbes1989">{{cite journal |last=Forbes |first=L. Scott |title=Prey Defences and Predator Handling Behaviour: The Dangerous Prey Hypothesis |journal=Oikos |volume=55 |issue=2 |year=1989|doi=10.2307/3565418 |pages=155–158 |jstor=3565418 |bibcode=1989Oikos..55..155F }}</ref>

===Solitary versus social predation===
{{See also|Cooperative hunting}}

In social predation, a group of predators cooperates to kill prey. This makes it possible to kill creatures larger than those they could overpower singly; for example, [[hyena]]s, and [[Wolf|wolves]] collaborate to catch and kill herbivores as large as buffalo, and lions even hunt elephants.<ref name="LangFarine2017">{{cite journal |last1=Lang |first1=Stephen D. J. |last2=Farine |first2=Damien R. |title=A multidimensional framework for studying social predation strategies |journal=Nature Ecology & Evolution |volume=1 |issue=9 |year=2017 |pages=1230–1239 |doi=10.1038/s41559-017-0245-0 |pmid=29046557 |bibcode=2017NatEE...1.1230L |url=https://ora.ox.ac.uk/objects/uuid:379eff38-903d-45d9-a19b-ccc01a114e9e }}</ref><ref name="MacNulty2014">{{cite journal |last1=MacNulty |first1=Daniel R. |last2=Tallian |first2=Aimee |last3=Stahler |first3=Daniel R. |last4=Smith |first4=Douglas W. | editor-last=Sueur | editor-first=Cédric |title=Influence of Group Size on the Success of Wolves Hunting Bison |journal=PLOS ONE|volume=9 |issue=11 | date=12 November 2014 |doi=10.1371/journal.pone.0112884 |pmid=25389760 |pmc=4229308 | article-number=e112884|bibcode=2014PLoSO...9k2884M |doi-access=free }}</ref><ref>{{cite journal |last1=Power |first1=R. John |last2=Shem Compion |first2=R.X. |title=Lion Predation on Elephants in the Savuti, Chobe National Park, Botswana |journal=African Zoology |date=April 2009 |volume=44 |issue=1 |pages=36–44 |doi=10.3377/004.044.0104 }}</ref> It can also make prey more readily available through strategies like flushing of prey and herding it into a smaller area. For example, when mixed flocks of birds forage, the birds in front flush out insects that are caught by the birds behind. [[Spinner dolphin]]s form a circle around a school of fish and move inwards, concentrating the fish by a factor of 200.<ref name=Beauchamp2012>{{harvnb|Beauchamp|2012|pages=7–12}}</ref> By hunting socially [[Common chimpanzee|chimpanzee]]s can catch [[colobus monkey]]s that would readily escape an individual hunter, while cooperating [[Harris hawk]]s can trap rabbits.<ref name="LangFarine2017"/><ref name=Dawson1988>{{cite thesis |last1=Dawson |first1=James W. |title=The cooperative breeding system of the Harris' Hawk in Arizona |hdl=10150/276864 |date=1988 |degree=Masters }}{{pn|date=July 2024}}</ref>

[[File:Journal.pone.0112884.g001 a.png|thumb|upright=2.3|center|[[Wolf|Wolves]], [[social predator]]s, cooperate to hunt and kill [[bison]].]]

Predators of different species sometimes cooperate to catch prey. In [[coral reefs]], when fish such as the [[grouper]] and [[coral trout]] spot prey that is inaccessible to them, they signal to [[Giant moray|giant moray eels]], [[Humphead wrasse|Napoleon wrasses]] or [[octopus]]es. These predators are able to access small crevices and flush out the prey.<ref name=Vail2013>{{cite journal |last1=Vail |first1=Alexander L. |last2=Manica |first2=Andrea |last3=Bshary |first3=Redouan |title=Referential gestures in fish collaborative hunting |journal=Nature Communications |date=23 April 2013 |volume=4 |issue=1 |page=1765 |doi=10.1038/ncomms2781|pmid=23612306 |doi-access=free |bibcode=2013NatCo...4.1765V }}</ref><ref>{{cite news |last1=Yong |first1=Ed |title=Groupers Use Gestures to Recruit Morays For Hunting Team-Ups |url=https://www.nationalgeographic.com/science/phenomena/2013/04/24/groupers-use-gestures-to-recruit-morays-for-hunting-team-ups/ |archive-url=https://web.archive.org/web/20180917215312/https://www.nationalgeographic.com/science/phenomena/2013/04/24/groupers-use-gestures-to-recruit-morays-for-hunting-team-ups/ |archive-date=17 September 2018 |access-date=17 September 2018 |work=National Geographic |date=24 April 2013}}</ref> [[Killer whale]]s have been known to help whalers hunt [[baleen whale]]s.<ref name=eden>{{cite AV media|people=Toft, Klaus (Producer) |date=2007 |url=http://shop.abc.net.au/browse/product.asp?productid=743192 |title=Killers in Eden (DVD documentary) |publisher=[[Australian Broadcasting Corporation]] |archive-url=https://web.archive.org/web/20090812231151/http://shop.abc.net.au/browse/product.asp?productid=743192 |archive-date=12 August 2009 }} ISBN R-105732-9.</ref>

Social hunting allows predators to tackle a wider range of prey, but at the risk of competition for the captured food. Solitary predators have more chance of eating what they catch, at the price of increased expenditure of energy to catch it, and increased risk that the prey will escape.<ref name="BryceWilmers2017"/><ref name="MajerHolm2018">{{cite journal |last1=Majer |first1=Marija |last2=Holm |first2=Christina |last3=Lubin |first3=Yael |last4=Bilde |first4=Trine |title=Cooperative foraging expands dietary niche but does not offset intra-group competition for resources in social spiders|journal=Scientific Reports |volume=8 |issue=1 |page=11828 |year=2018 |doi=10.1038/s41598-018-30199-x|pmid=30087391 |pmc=6081395 |bibcode=2018NatSR...811828M }}</ref> Ambush predators are often solitary to reduce the risk of becoming prey themselves.<ref>{{cite web |title=Ambush Predators |url=https://sibleynaturecenter.org/photo-essays/ambush-predators |website=Sibley Nature Center |access-date=17 September 2018 |language=en |archive-date=2 August 2021 |archive-url=https://web.archive.org/web/20210802002919/https://sibleynaturecenter.org/photo-essays/ambush-predators }}</ref> Of 245 terrestrial members of the [[Carnivora]] (the group that includes the cats, dogs, and bears), 177 are solitary; and 35 of the 37 [[Felidae|wild cats]] are solitary,<ref>{{cite journal |last1=Elbroch |first1=L. Mark |last2=Quigley |first2=Howard |title=Social interactions in a solitary carnivore |journal=Current Zoology |volume=63 |issue=4 |date=10 July 2016 |pages=357–362 |doi=10.1093/cz/zow080|pmid=29491995 |pmc=5804185 }}</ref> including the cougar and cheetah.<ref name="BryceWilmers2017">{{cite journal |last1=Bryce |first1=Caleb M. |last2=Wilmers |first2=Christopher C. |last3=Williams |first3=Terrie M. |title=Energetics and evasion dynamics of large predators and prey: pumas vs. hounds |journal=PeerJ |volume=5 |year=2017 |article-number=e3701 |doi=10.7717/peerj.3701|pmid=28828280 |pmc=5563439 |doi-access=free }}</ref><ref name=LaffertyKuris2002/> However, the solitary cougar does allow other cougars to share in a kill,<ref>{{cite news |last1=Quenqua |first1=Douglas |title=Solitary Pumas Turn Out to Be Mountain Lions Who Lunch |url=https://www.nytimes.com/2017/10/11/science/pumas-solitary-social.html?hpw&rref=science&action=click&pgtype=Homepage&module=well-region&region=bottom-well&WT.nav=bottom-well |access-date=17 September 2018 |work=The New York Times |date=11 October 2017 |language=en}}</ref> and the [[coyote]] can be either solitary or social.<ref>{{cite book |last1=Flores |first1=Dan |title=Coyote America: a natural and supernatural history |date=2016 |publisher=Basic Books |isbn=978-0-465-05299-8}}</ref> Other solitary predators include the northern pike,<ref name="StowNyqvist2012">{{cite journal |last1=Stow |first1=Adam |last2=Nyqvist |first2=Marina J. |last3=Gozlan |first3=Rodolphe E. |last4=Cucherousset |first4=Julien |last5=Britton |first5=J. Robert |title=Behavioural Syndrome in a Solitary Predator Is Independent of Body Size and Growth Rate |journal=PLOS ONE|volume=7 |issue=2 |year=2012 |article-number=e31619 |doi=10.1371/journal.pone.0031619|pmid=22363687 |pmc=3282768 |bibcode=2012PLoSO...731619N |doi-access=free }}</ref> [[wolf spider]]s and all the thousands of species of [[solitary wasp]]s among arthropods,<ref>{{cite web |title=How do Spiders Hunt? |url=https://www.amnh.org/explore/news-blogs/on-exhibit-posts/how-do-spiders-hunt/ |publisher=American Museum of Natural History |access-date=5 September 2018 |date=25 August 2014}}</ref><ref name="WeselohHare2009">{{cite encyclopedia |last1=Weseloh |first1=Ronald M. |title=Encyclopedia of Insects |last2=Hare |first2=J. Daniel |edition=Second |year=2009 |pages=837–839 |doi=10.1016/B978-0-12-374144-8.00219-8|isbn=978-0-12-374144-8 |chapter=Predation/Predatory Insects }}</ref> and many [[microorganism]]s and [[zooplankton]].<ref name=VelicerMendes-Soares2007>{{cite journal |last1=Velicer |first1=Gregory J. |last2=Mendes-Soares |first2=Helena |title=Bacterial predators |journal=Cell |date=2007 |volume=19 |issue=2 |pages=R55–R56 |doi=10.1016/j.cub.2008.10.043 |pmid=19174136 |s2cid=5432036 |doi-access=free }}</ref>

==Specialization==

===Physical adaptations===

Under the pressure of [[natural selection]], predators have evolved a variety of physical [[adaptation]]s for detecting, catching, killing, and digesting prey. These include speed, agility, stealth, sharp senses, claws, teeth, filters, and suitable digestive systems.<ref>{{cite web |last1=Bar-Yam |title=Predator-Prey Relationships |url=http://necsi.edu/projects/evolution/co-evolution/pred-prey/co-evolution_predator.html |publisher=New England Complex Systems Institute |access-date=7 September 2018}}</ref>

For [[Prey detection|detecting prey]], predators have well-developed [[eye|vision]], [[olfaction|smell]], or [[hearing]].<ref name=Stevens2010/> Predators as diverse as [[owl]]s and [[jumping spider]]s have forward-facing eyes, providing accurate [[binocular vision]] over a relatively narrow field of view, whereas prey animals often have less acute all-round vision. Animals such as foxes can smell their prey even when it is concealed under {{convert|2|ft|cm|-1}} of snow or earth. Many predators have acute hearing, and some such as [[animal echolocation|echolocating]] [[bat]]s hunt exclusively by active or passive use of sound.<ref name="RSM2012">{{cite web |title=Predator & Prey: Adaptations |url= https://royalsaskmuseum.ca/pub/Lesson%20Plans/Resources/Predator%20and%20Prey%20Adaptations.pdf |publisher= Royal Saskatchewan Museum |access-date= 19 April 2018 |date= 2012 |archive-date= 3 April 2018 |archive-url= https://web.archive.org/web/20180403131659/http://royalsaskmuseum.ca/pub/Lesson%20Plans/Resources/Predator%20and%20Prey%20Adaptations.pdf }}</ref>

Predators including [[big cat]]s, [[bird of prey|birds of prey]], and ants share powerful jaws, sharp teeth, or claws which they use to seize and kill their prey. Some predators such as [[snake]]s and fish-eating birds like [[herons]] and [[cormorants]] swallow their prey whole; some snakes can unhinge their jaws to allow them to swallow large prey, while fish-eating birds have long spear-like beaks that they use to stab and grip fast-moving and slippery prey.<ref name="RSM2012"/> Fish and other predators have developed the ability to crush or open the armoured shells of molluscs.<ref>{{cite book |last=Vermeij |first=Geerat J. |title=Evolution and Escalation: An Ecological History of Life |url=https://books.google.com/books?id=M3pCQ6ks5PEC&pg=PR11 |year=1993 |publisher=[[Princeton University Press]] |isbn=978-0-691-00080-0 |pages=11 and passim}}</ref>

Many predators are powerfully built and can catch and kill animals larger than themselves; this applies as much to small predators such as [[ant]]s and [[shrew]]s as to big and visibly muscular carnivores like the [[cougar]] and [[lion]].<ref name="RSM2012"/><ref name=LaffertyKuris2002>{{cite journal |last1=Lafferty |first1=K. D. |last2=Kuris |first2=A. M. |date=2002 |title=Trophic strategies, animal diversity and body size |journal=Trends Ecol. Evol. |volume=17 |issue=11 |pages=507–513 |doi=10.1016/s0169-5347(02)02615-0}}</ref><ref name=Getz2011>{{cite journal |last=Getz |first=W. M. |title=Biomass transformation webs provide a unified approach to consumer-resource modelling |journal=Ecology Letters |volume=14 |issue=2 |pages=113–24 |year=2011 |pmid=21199247 |pmc=3032891 |doi=10.1111/j.1461-0248.2010.01566.x |bibcode=2011EcolL..14..113G }}</ref>

<gallery mode="packed">
File:Ursus arctos 01 MWNH 145 (cropped).JPG|Skull of [[brown bear]] has large pointed [[canine tooth|canines]] for killing prey, and self-sharpening [[carnassial]] teeth at rear for cutting flesh with a scissor-like action
File:Myrmecia pilosula specimen mandibles.jpg|Large [[compound eye]]s, sensitive [[antenna (biology)|antennae]], and powerful jaws ([[mandible (insect)|mandibles]]) of [[Myrmecia pilosula|jack jumper ant]]
File:Crab spider seizes field digger wasp.jpg|[[Crab spider]], an [[ambush predator]] with forward-facing eyes, catching another predator, a [[Mellinus arvensis|field digger wasp]]
File:Hawk eating prey (cropped).jpg|[[Red-tailed hawk]] uses sharp hooked claws and beak to kill and tear up its prey
File:GreatBlueHeronTampaFL.JPG|Specialist: a [[great blue heron]] with a speared fish
File:MNP Python at Moyer.jpg|[[Python molurus|Indian python]] unhinges its jaw to swallow large prey like this [[chital]]
</gallery>

===Diet and behaviour===
{{further|Generalist and specialist species}}

{{multiple image|align=right|image1=Peerj-297-fig-5 Platydemus manokwari.png|width1=200|caption1=''[[Platydemus manokwari]]'', a specialist [[flatworm]] predator of [[land snail]]s, attacking a snail|image2=Lioness vs Cape Buffalo (cropped).jpg|width2=212|caption2=Size-selective predation: a [[lion]]ess attacking a [[Cape buffalo]], over twice her weight. Lions can attack much larger prey, including elephants, but do so much less often.}}

Predators are often highly specialized in their diet and hunting behaviour; for example, the [[Eurasian lynx]] only hunts small [[ungulate]]s.<ref name="Sidorovich2011">{{cite book |last=Sidorovich |first=Vadim |title=Analysis of vertebrate predator-prey community: Studies within the European Forest zone in terrains with transitional mixed forest in Belarus |url=https://books.google.com/books?id=cHxwAwAAQBAJ&pg=PA426 |year=2011 |publisher=Tesey |isbn=978-985-463-456-2 |page=426}}</ref> Others such as [[leopard]]s are more opportunistic generalists, preying on at least 100 species.<ref name="Angelici2015">{{cite book |last=Angelici |first=Francesco M. |title=Problematic Wildlife: A Cross-Disciplinary Approach |url=https://books.google.com/books?id=S1c-CwAAQBAJ&pg=PA160 |year=2015 |publisher=Springer |isbn=978-3-319-22246-2 |page=160}}</ref><ref name=Hayward2006>{{cite journal |last1=Hayward |first1=M. W. |last2=Henschel |first2=P. |last3=O'Brien |first3=J. |last4=Hofmeyr |first4=M. |last5=Balme |first5=G. |last6=Kerley |first6=G.I.H. |title=Prey preferences of the leopard (''Panthera pardus'') |journal=Journal of Zoology |date=2006 |volume=270 |issue=2 |pages=298–313 |doi=10.1111/j.1469-7998.2006.00139.x }}</ref> The specialists may be highly adapted to capturing their preferred prey, whereas generalists may be better able to switch to other prey when a preferred target is scarce. When prey have a clumped (uneven) distribution, the optimal strategy for the predator is predicted to be more specialized as the prey are more conspicuous and can be found more quickly;<ref name=Pulliam1974>{{cite journal |last1=Pulliam |first1=H. Ronald |title=On the Theory of Optimal Diets |journal=The American Naturalist |volume=108 |issue=959 |year=1974 |pages=59–74 |doi=10.1086/282885|bibcode=1974ANat..108...59P |s2cid=8420787 }}</ref> this appears to be correct for predators of immobile prey, but is doubtful with mobile prey.<ref name="SihChristensen2001">{{cite journal |last1=Sih |first1=Andrew |last2=Christensen |first2=Bent |title=Optimal diet theory: when does it work, and when and why does it fail? |journal=Animal Behaviour|volume=61 |issue=2 |year=2001 |doi=10.1006/anbe.2000.1592 |pages=379–390| s2cid=44045919 }}</ref>

{{anchor|Size-selective predation}}
In size-selective predation, predators select prey of a certain size.<ref>{{cite journal |last=Sprules |first=W. Gary |title=Effects of Size-Selective Predation and Food Competition on High Altitude Zooplankton Communities |journal=Ecology |volume=53 |issue=3 |year=1972 |doi=10.2307/1934223 |jstor=1934223 |pages=375–386|bibcode=1972Ecol...53..375S }}</ref> Large prey may prove troublesome for a predator, while small prey might prove hard to find and in any case provide less of a reward. This has led to a correlation between the size of predators and their prey. Size may also act as a [[refuge (ecology)|refuge]] for large prey. For example, adult elephants are relatively safe from predation by lions, but juveniles are vulnerable.<ref name="Owen-SmithMills2008">{{cite journal |last1=Owen-Smith |first1=Norman |last2=Mills |first2=M. G. L. |title=Predator-prey size relationships in an African large-mammal food web |journal=Journal of Animal Ecology |volume=77 |issue=1 |year=2008 |doi=10.1111/j.1365-2656.2007.01314.x |pmid=18177336 |pages=173–183|bibcode=2008JAnEc..77..173O | hdl=2263/9023 |doi-access=free |hdl-access=free }}</ref>

=== Camouflage and mimicry ===
{{further|Camouflage|Aggressive mimicry}}

{{multiple image
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| image1 = Snow Leopard in Ladakh( Photo by Tashi Lonchay).jpg
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| caption1 = A [[camouflage]]d predator: [[snow leopard]] in [[Ladakh]]
| image2 = Striped anglerfish ( Antennarius striatus ).jpg
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| caption2 = [[Striated frogfish]] uses camouflage and [[aggressive mimicry]] in the form of a fishing rod-like [[Fin ray|lure]] on its head to attract prey.
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}}

Members of the [[Felidae|cat family]] such as the [[snow leopard]] (treeless highlands), [[tiger]] (grassy plains, reed swamps), [[ocelot]] (forest), [[fishing cat]] (waterside thickets), and [[lion]] (open plains) are camouflaged with coloration and [[disruptive pattern]]s suiting their habitats.<ref name=Cott12>{{harvnb|Cott|1940|pages=12–13}}</ref>

In aggressive mimicry, certain predators, including insects and fishes, make use of coloration and behaviour to attract prey. Female ''[[Photuris (genus)|Photuris]]'' [[Firefly|fireflies]], for example, copy the light signals of other species, thereby attracting male fireflies, which they capture and eat.<ref>{{cite journal | author=Lloyd J. E. |year=1965 |title=Aggressive Mimicry in Photuris: Firefly Femmes Fatales |  journal=[[Science (journal)|Science]] |volume=149 |issue=3684|pages=653–654 |doi=10.1126/science.149.3684.653 |pmid=17747574 |bibcode=1965Sci...149..653L | s2cid=39386614 }}</ref> [[Flower mantis]]es are ambush predators; camouflaged as flowers, such as [[orchid]]s, they attract prey and seize it when it is close enough.<ref>{{cite book |last=Forbes |first=Peter |year=2009 |title=Dazzled and Deceived: Mimicry and Camouflage |publisher=Yale University Press |isbn=978-0-300-17896-8 |page=134|title-link=Dazzled and Deceived: Mimicry and Camouflage }}</ref> [[Frogfish]]es are extremely well camouflaged, and actively lure their prey to approach using an [[Fin ray|esca]], a bait on the end of a rod-like appendage on the head, which they wave gently to mimic a small animal, gulping the prey in an extremely rapid movement when it is within range.<ref>{{cite web |last=Bester |first=Cathleen |date=2 May 2025 |title=Striated frogfish |url= https://www.floridamuseum.ufl.edu/discover-fish/species-profiles/striated-frogfish/ |website=Florida Museum |publisher=University of Florida |access-date=12 July 2025}}</ref>

===Venom===
{{further|Venom|Evolution of snake venom}}

Many smaller predators such as the [[box jellyfish]] use [[venom]] to subdue their prey,<ref>{{cite book |title=Invertebrate Zoology |edition=7th |last1=Ruppert |first1=Edward E. |last2=Fox |first2=Richard S. |last3=Barnes |first3=Robert D. |year=2004 |publisher=Cengage Learning |isbn=978-81-315-0104-7 |pages=153–154 }}</ref> and venom can also aid in digestion (as is the case for [[rattlesnake]]s and some [[spider]]s).<ref>{{cite book |author=Cetaruk, Edward W. |chapter=Rattlesnakes and Other Crotalids |editor=Brent, Jeffrey |title=Critical care toxicology: diagnosis and management of the critically poisoned patient |publisher=Elsevier Health Sciences |year=2005 |isbn=978-0-8151-4387-1 |page=1075 |chapter-url=https://books.google.com/books?id=WuA4LsWXXWEC&pg=PA1075}}</ref><ref name="Barceloux2008">{{cite book |last=Barceloux |first=Donald G. |title=Medical Toxicology of Natural Substances: Foods, Fungi, Medicinal Herbs, Plants, and Venomous Animals |url=https://books.google.com/books?id=CpqzhHc072AC&pg=PA1028 |year=2008 |publisher=Wiley |isbn=978-0-470-33557-4 |page=1028}}</ref> The [[marbled sea snake]] that has adapted to egg predation has atrophied venom glands, and the gene for its three finger toxin contains a [[mutation]] (the deletion of two [[nucleotide]]s) that inactives it. These changes are explained by the fact that its prey does not need to be subdued.<ref name="Li Fry Kini 2005 pp. 81–89">{{cite journal |last1=Li |first1=Min |last2=Fry |first2=B.G. |last3=Kini |first3=R. Manjunatha |title=Eggs-Only Diet: Its Implications for the Toxin Profile Changes and Ecology of the Marbled Sea Snake (Aipysurus eydouxii) |journal=Journal of Molecular Evolution |volume=60 |issue=1 |year=2005 |doi=10.1007/s00239-004-0138-0 |pmid=15696370 |pages=81–89 |bibcode=2005JMolE..60...81L| s2cid=17572816 }}</ref>

===Electric fields===
[[File:Elektroplax_Rochen.png|thumb|An [[electric ray]] ([[Torpediniformes]]) showing location of electric organ and electrocytes stacked within it]]
{{further|Electroreception and electrogenesis|Electric organ (fish)}}

Several groups of predatory fish have the ability to detect, track, and sometimes, as in the [[electric ray]], to incapacitate their prey by [[Electroreception and electrogenesis|sensing and generating electric fields]].<ref>{{cite journal |author1= Castello, M. E. |author2=A. Rodriguez-Cattaneo |author3=P. A. Aguilera |author4=L. Iribarne |author5=A. C. Pereira |author6=A. A. Caputi |title=Waveform generation in the weakly electric fish Gymnotus coropinae (Hoedeman): the electric organ and the electric organ discharge |journal=Journal of Experimental Biology |volume=212 |year=2009 |pages=1351–1364 |doi=10.1242/jeb.022566 |issue=9 |pmid=19376956 |doi-access=free |bibcode=2009JExpB.212.1351C }}</ref><ref name="Feulner, P. G., M. Plath, J. Engelmann, F. Kirschbaum, R. Tiedemann 2009 225–228">{{cite journal |author1= Feulner, P. G. |author2=M. Plath |author3=J. Engelmann |author4=F. Kirschbaum |author5=R. Tiedemann |title=Electrifying love: electric fish use species-specific discharge for mate recognition |journal=Biology Letters |volume=5 |issue=2 |year=2009 |pages=225–228 |doi=10.1098/rsbl.2008.0566|pmid=19033131 |pmc= 2665802 }}</ref><ref name="Catania2015">{{cite journal |last= Catania |first= Kenneth C. |title= Electric eels use high-voltage to track fast-moving prey |journal= Nature Communications |volume=6 |issue=1 |year=2015 |article-number=8638 |doi=10.1038/ncomms9638 |pmid=26485580 |pmc=4667699 |bibcode=2015NatCo...6.8638C}}</ref> The electric organ is derived from modified nerve or muscle tissue.<ref name=Kramer1996>{{cite book |doi=10.5283/epub.2108 |date=1996 |last1=Kramer |first1=Bernd |title=Electroreception and communication in fishes |volume=42 |publisher=Universität Regensburg |isbn=978-3-437-25038-5 }}</ref>

===Physiology===

Physiological adaptations to predation include the ability of predatory bacteria to digest the complex [[peptidoglycan]] polymer from the [[cell wall]]s of the bacteria that they prey upon.<ref name="JurkevitchDavidov2006"/> Carnivorous vertebrates of all five major classes (fishes, amphibians, reptiles, birds, and mammals) have lower relative rates of sugar to [[amino acid]] transport than either herbivores or omnivores, presumably because they acquire plenty of amino acids from the animal [[protein]]s in their diet.<ref name="KarasovDiamond1988">{{cite journal |last1=Karasov |first1=William H. |last2=Diamond |first2=Jared M. |title=Interplay between Physiology and Ecology in Digestion |journal=BioScience |volume=38 |issue=9 |year=1988 |doi=10.2307/1310825 |pages=602–611 |jstor=1310825}}</ref>

== Antipredator adaptations ==
{{main|Antipredator adaptation}}

{{multiple image |align=right|image1=Bristol.zoo.dead.leaf.mantis.arp.jpg |width1=190 |caption1=[[Dead leaf mantis]]'s [[camouflage]] makes it less visible to both predators and prey. |image2=Syrphid hoverfly wasp mimic.jpg |width2=157 |caption2=[[Syrphidae|Syrphid hoverfly]] misdirects predators by [[mimicry|mimicking]] a [[wasp]], but has no [[stinger|sting]].}}

To counter predation, prey have evolved defences for use at each stage of an attack.<ref>{{harvnb|Ruxton|Sherratt|Speed|2004|pages=vii–xii}}</ref><ref name=Stevens2010/> They can try to avoid detection,<ref name=CaroCh3>{{harvnb|Caro|2005|pages=67–114}}</ref> such as by using [[camouflage]] and [[mimicry]].<ref name="Merilaita Scott-Samuel Cuthill p=20160341">{{cite journal |last1=Merilaita |first1=Sami |last2=Scott-Samuel |first2=Nicholas E. |last3=Cuthill |first3=Innes C. | author3-link=Innes Cuthill |title=How camouflage works |journal=Philosophical Transactions of the Royal Society B: Biological Sciences |volume=372 |issue=1724 | date=22 May 2017 |doi=10.1098/rstb.2016.0341 |pmid=28533458 |pmc=5444062 }}</ref> They can detect predators<ref name=Caro13>{{harvnb|Caro|2005|pages=13–15}}</ref> and warn others of their presence.<ref>{{cite journal |last1=Bergstrom |first1=C. T. |last2=Lachmann |first2=M. |year=2001 |title=Alarm calls as costly signals of antipredator vigilance: the watchful babbler game |journal=Animal Behaviour |volume=61 |issue=3 |pages=535–543 |doi=10.1006/anbe.2000.1636 |s2cid=2295026 }}</ref><ref>{{cite journal |last=Getty |first=T. |date=2002 |title=The discriminating babbler meets the optimal diet hawk |journal=Animal Behaviour |volume=63 |issue=2 |pages=397–402 |doi=10.1006/anbe.2001.1890|s2cid=53164940 }}</ref>
If detected, they can try to avoid being the target of an attack, for example, by [[Aposematism|signalling that they are toxic or unpalatable]],<ref>{{harvnb|Cott|1940|pages=241–307}}</ref><ref name=Bowers1985>{{cite journal |last1=Bowers |first1=M. D. |last2=Brown |first2=Irene L. |last3=Wheye |first3=Darryl |year=1985 |title=Bird Predation as a Selective Agent in a Butterfly Population |journal=Evolution |volume=39 |issue=1|pages=93–103 |doi=10.1111/j.1558-5646.1985.tb04082.x |pmid=28563638 | s2cid=12031679 |doi-access=free }}</ref><ref name=Berenbaum1995>{{cite journal |last=Berenbaum |first=M. R. |title=The chemistry of defense: theory and practice |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=92 |issue=1 | date=3 January 1995 |pmid=7816816 |pmc=42807 |pages=2–8|doi=10.1073/pnas.92.1.2 |bibcode=1995PNAS...92....2B |doi-access=free }}</ref> by signalling that a chase would be unprofitable,<ref name=Ruxton70>{{harvnb|Ruxton|Sherratt|Speed|2004|pages=70–81}}</ref><ref name=Caro1986>{{harvnb|Caro|2005|pages=663–684}}</ref> or by forming groups.<ref name=BeauchampCh3>{{harvnb|Beauchamp|2012|pages=83–88}}</ref><ref>{{cite book |last1=Krause |first1=Jens |last2=Ruxton |first2=Graeme D. |title=Living in groups |publisher=Oxford University Press |isbn=978-0-19-850818-2 |pages=13–15|date=10 October 2002 }}</ref> If they become a target, they can try to fend off the attack with defences such as armour, [[Spine (zoology)|quills]], unpalatability, or mobbing;<ref name=RuxtonSherrattSpeed2004>{{harvnb|Ruxton|Sherratt|Speed|2004|pages=54–55}}</ref><ref name="Dominey">{{cite journal |last=Dominey |first=Wallace J. |year=1983 |title=Mobbing in Colonially Nesting Fishes, Especially the Bluegill, ''Lepomis macrochirus'' |journal=Copeia |volume=1983 |issue=4 |pages=1086–1088 |doi=10.2307/1445113 |jstor=1445113}}</ref><ref name=Brodie2009>{{cite journal |last=Brodie |first=Edmund D. |title=Toxins and venoms |journal=Current Biology |volume=19 |issue=20 |pages=R931–R935 |date=3 November 2009 |doi=10.1016/j.cub.2009.08.011 |pmid=19889364 |s2cid=9744565 |doi-access=free |bibcode=2009CBio...19.R931B }}</ref> and they can often escape an attack in progress by [[deimatic behaviour|startling]] the predator,<ref>{{harvnb|Cott|1940|pages=368–389}}</ref><ref name=Merilaita2011>{{Cite journal |volume=22 |issue=6 |pages=1326–1331 | date=26 July 2011 |last1=Merilaita |first1=Sami |title=Number of eyespots and their intimidating effect on naïve predators in the peacock butterfly |journal=Behavioral Ecology |last2=Vallin |first2=Adrian  |author3=Kodandaramaiah, Ullasa |author4=Dimitrova, Marina |author5=Ruuskanen, Suvi |author6=Laaksonen, Toni |display-authors=3 |doi=10.1093/beheco/arr135 |doi-access=free }}</ref><ref name=Springer>{{cite book |doi=10.1007/978-1-4020-6359-6_863 |chapter=Deimatic Behavior |title=Encyclopedia of Entomology |date=2008 |last1=Cumming |first1=Jeffrey M. |last2=Sinclair |first2=Bradley J. |last3=Triplehorn |first3=Charles A. |last4=Aldryhim |first4=Yousif |last5=Galante |first5=Eduardo |last6=Marcos-Garcia |first6=Ma Angeles |last7=Edmunds |first7=Malcolm |last8=Edmunds |first8=Malcolm |last9=Lounibos |first9=L. Phillip |last10=Frank |first10=J. Howard |last11=Showler |first11=Allan T. |last12=Yu |first12=Simon J. |last13=Capinera |first13=John L. |last14=Heppner |first14=John B. |last15=Philogène |first15=Bernard J. R. |last16=Lapointe |first16=Stephen L. |last17=Capinera |first17=John L. |last18=Capinera |first18=John L. |last19=Nayar |first19=Jai K. |last20=Goettel |first20=Mark S. |last21=Nation |first21=James L. |last22=Heppner |first22=John B. |last23=Negron |first23=Jose F. |last24=Heppner |first24=John B. |last25=Kondratieff |first25=Boris C. |last26=Schöning |first26=Caspar |last27=Stewart |first27=Kenneth W. |last28=Aldryhim |first28=Yousif |last29=Heppner |first29=John B. |last30=Hangay |first30=George |pages=1173–1174 |isbn=978-1-4020-6242-1 }}</ref> [[Thanatosis|playing dead]], [[autotomy|shedding body parts]] such as tails, or simply fleeing.<ref>{{harvnb|Caro|2005|pages=v–xi, 4–5}}</ref><ref name=CaroCh12>{{harvnb|Caro|2005|page=413–414}}</ref>

== Coevolution ==
{{further|Coevolution}}
[[File:Big-eared-townsend-fledermaus.jpg|thumb|left|Bats use [[Animal echolocation|echolocation]] to hunt moths at night.]]

Predators and prey are natural enemies, and many of their adaptations seem designed to counter each other. For example, bats have sophisticated [[Animal echolocation|echolocation]] systems to detect insects and other prey, and insects have developed a variety of defences including the ability to hear the echolocation calls.<ref>{{harvnb|Jacobs|Bastian|2017|page=4}}</ref><ref>{{cite book |last1=Barbosa |first1=Pedro |last2=Castellanos |first2=Ignacio |title=Ecology of predator-prey interactions |date=2005 |publisher=Oxford University Press |isbn=978-0-19-987454-5|url=https://archive.org/details/ecologypredatorp00barb|url-access=limited |page=[https://archive.org/details/ecologypredatorp00barb/page/n96 78]}}</ref> Many pursuit predators that run on land, such as wolves, have evolved long limbs in response to the increased speed of their prey.<ref name=Janis>{{cite journal |last1=Janis |first1=C. M. |last2=Wilhelm |first2=P. B. |year=1993 |title=Were there mammalian pursuit predators in the Tertiary? Dances with wolf avatars |journal=Journal of Mammalian Evolution |volume=1 |issue=2|pages=103–125 |doi=10.1007/bf01041590| s2cid=22739360 }}</ref> Their adaptations have been characterized as an [[evolutionary arms race]], an example of the [[coevolution]] of two species.<ref name=DawKrebs>{{cite journal |author1=Dawkins, Richard |author1-link=Richard Dawkins |author2=Krebs, J. R. |author2-link=John Krebs, Baron Krebs |year=1979 |title=Arms races between and within species |journal=Proceedings of the Royal Society B: Biological Sciences |volume=205 |issue=1161 |pages=489–511|doi=10.1098/rspb.1979.0081 |pmid=42057 |bibcode=1979RSPSB.205..489D |s2cid=9695900 }}</ref> In a [[gene centered view of evolution]], the genes of predator and prey can be thought of as [[Competition (biology)|competing]] for the prey's body.<ref name=DawKrebs/> However, the "life-dinner" principle of Dawkins and Krebs predicts that this arms race is asymmetric: if a predator fails to catch its prey, it loses its dinner, while if it succeeds, the prey loses its life.<ref name=DawKrebs/>

[[File:Micrurus fulviusHolbrookV3P10AA.jpg|thumb|upright=0.7|[[Micrurus fulvius|Eastern coral snake]], itself a predator, is venomous enough to kill predators that attack it, so when they avoid it, this behaviour must be inherited, not learnt.]]

The metaphor of an arms race implies ever-escalating advances in attack and defence. However, these adaptations come with a cost; for instance, longer legs have an increased risk of breaking,<ref name=Abrams>{{cite journal |last1=Abrams |first1=Peter A. |title=Adaptive responses of predators to prey and prey to predators: The failure of the arms-race analogy |journal=Evolution |date=November 1986 |volume=40 |issue=6 |pages=1229–1247 |doi=10.1111/j.1558-5646.1986.tb05747.x|pmid=28563514 |s2cid=27317468 |doi-access=free }}</ref> while the specialized tongue of the chameleon, with its ability to act like a projectile, is useless for lapping water, so the chameleon must drink dew off vegetation.<ref name=Brodie>{{cite journal |last1=Brodie |first1=Edmund D. |title=Predator-Prey Arms Races |journal=BioScience |date=July 1999 |volume=49 |issue=7 |pages=557–568 |doi=10.2307/1313476|jstor=1313476 |doi-access=free }}</ref>

The "life-dinner" principle has been criticized on multiple grounds. The extent of the asymmetry in natural selection depends in part on the heritability of the adaptive traits.<ref name=Brodie/> Also, if a predator loses enough dinners, it too will lose its life.<ref name=Abrams/><ref name=Brodie/> On the other hand, the fitness cost of a given lost dinner is unpredictable, as the predator may quickly find better prey. In addition, most predators are generalists, which reduces the impact of a given prey adaption on a predator. Since specialization is caused by predator-prey coevolution, the rarity of specialists may imply that predator-prey arms races are rare.<ref name=Brodie/>

It is difficult to determine whether given adaptations are truly the result of coevolution, where a prey adaptation gives rise to a predator adaptation that is countered by further adaptation in the prey. An alternative explanation is ''escalation'', where predators are adapting to competitors, their own predators or dangerous prey.<ref>{{cite journal |last1=Vermeij |first1=G J |title=The Evolutionary Interaction Among Species: Selection, Escalation, and Coevolution |journal=Annual Review of Ecology and Systematics |date=November 1994 |volume=25 |issue=1 |pages=219–236 |doi=10.1146/annurev.es.25.110194.001251|bibcode=1994AnRES..25..219V }}</ref> Apparent adaptations to predation may also have arisen for other reasons and then been co-opted for attack or defence. In some of the insects preyed on by bats, hearing evolved before bats appeared and was used to hear signals used for territorial defence and mating.<ref>{{harvnb|Jacobs|Bastian|2017|page=8}}</ref> Their hearing evolved in response to bat predation, but the only clear example of reciprocal adaptation in bats is stealth echolocation.<ref>{{harvnb|Jacobs|Bastian|2017|page=107}}</ref>

A more symmetric arms race may occur when the prey are dangerous, having spines, quills, toxins or venom that can harm the predator. The predator can respond with avoidance, which in turn drives the evolution of mimicry. Avoidance is not necessarily an evolutionary response as it is generally learned from bad experiences with prey. However, when the prey is capable of killing the predator (as can a [[coral snake]] with its venom), there is no opportunity for learning and avoidance must be inherited. Predators can also respond to dangerous prey with counter-adaptations. In western North America, the [[common garter snake]] has developed a resistance to the toxin in the skin of the [[rough-skinned newt]].<ref name=Brodie/>

==Role in ecosystems==

Predators affect their ecosystems not only directly by eating their own prey, but by indirect means such as reducing predation by other species, or altering the foraging behaviour of a herbivore, as with the biodiversity effect of wolves on riverside vegetation or sea otters on kelp forests. This may explain population dynamics effects such as the cycles observed in lynx and snowshoe hares.<ref>{{cite journal |last1=Sheriff |first1=Michael J. |last2=Peacor |first2=Scott D. |last3=Hawlena |first3=Dror |last4=Thaker |first4=Maria |last5=Gaillard |first5=Jean-Michel |title=Non-Consumptive Predator Effects on Prey Population Size: A Dearth of Evidence |journal=Journal of Animal Ecology |volume=89 |issue=6 |year=2020 |pages=1302–1316 |doi=10.1111/1365-2656.13213 |pmid=32215909 |bibcode=2020JAnEc..89.1302S |doi-access=free }}</ref><ref name="Preisser Bolnick 2005">{{cite journal |last1=Preisser |first1=Evan L. |last2=Bolnick |first2=Daniel I. |last3=Benard |first3=Michael F. |title=Scared to Death? The Effects of Intimidation and Consumption in Predator–Prey Interactions |journal=Ecology |volume=86 |issue=2 |year=2005 |pages=501–509 |issn=0012-9658 |doi=10.1890/04-0719|bibcode=2005Ecol...86..501P }}</ref><ref name=classics/>

=== Trophic level ===

{{further|Trophic level|Apex predator|Energy flow (ecology)}}

One way of classifying predators is by [[trophic level]]. [[Carnivore]]s that feed on [[herbivore]]s are secondary consumers; their predators are tertiary consumers, and so forth.<ref name="Lindeman1942">{{cite journal |last1=Lindeman |first1=Raymond L. |title=The Trophic-Dynamic Aspect of Ecology |journal=Ecology |volume=23 |issue=4 |year=1942 |pages=399–417 |doi=10.2307/1930126 |jstor=1930126 |bibcode=1942Ecol...23..399L }}</ref> At the top of this [[food chain]] are [[apex predator]]s such as [[lion]]s.<ref name="OrdizBischofSwenson2013">{{cite journal |last1=Ordiz |first1=Andrés |last2=Bischof |first2=Richard |last3=Swenson |first3=Jon E. |title=Saving large carnivores, but losing the apex predator? |journal=Biological Conservation |volume=168 |year=2013 |doi=10.1016/j.biocon.2013.09.024 |pages=128–133|doi-access=free |bibcode=2013BCons.168..128O | hdl=11250/2492589 | hdl-access=free }}</ref> Many predators however eat from multiple levels of the food chain; a carnivore may eat both secondary and tertiary consumers.<ref>{{Cite journal  |last1=Pimm |first1=S. L. |last2=Lawton |first2=J. H. |year=1978 |title=On feeding on more than one trophic level |journal=Nature |volume=275 |issue=5680 |pages=542–544 |doi=10.1038/275542a0 |bibcode=1978Natur.275..542P| s2cid=4161183 }}</ref> This means that many predators must contend with [[intraguild predation]], where other predators kill and eat them. For example, [[coyote]]s compete with and sometimes kill [[gray fox]]es and [[bobcat]]s.<ref>{{cite journal |last1=Fedriani |first1=Jose M. |last2=Fuller |first2=Todd K. |last3=Sauvajot |first3=Raymond M. |last4=York |first4=Eric C. |title=Competition and intraguild predation among three sympatric carnivores |journal=Oecologia |date=October 2000 |volume=125 |issue=2 |pages=258–270 |doi=10.1007/s004420000448 |pmid=24595837 |bibcode=2000Oecol.125..258F |hdl=10261/54628 |s2cid=24289407 |url=https://scholarworks.umass.edu/nrc_faculty_pubs/110 | hdl-access=free }}</ref>

Trophic transfer efficiency measures how effectively energy is passed up to higher trophic levels by predation. Each transfer decreases the available energy due to heat, waste, and the natural [[Metabolism|metabolic processes]] that occur as predators consume their prey. The result is that only about 10% of the energy at a trophic level is transferred to the next level. This limits the number of trophic levels that an individual ecosystem is capable of supporting.<ref>{{Cite web |date=18 February 2023 |title=Energy transfer in ecosystems |url=https://education.nationalgeographic.org/resource/energy-transfer-ecosystems/ |access-date=18 February 2023 |website=National Geographic}}</ref>

=== Biodiversity maintained by apex predation ===

{{further|Keystone species}}

Predators may increase the [[biodiversity]] of communities by preventing a single species from becoming dominant. Such predators are known as [[keystone species]] and may have a profound influence on the balance of organisms in a particular [[ecosystem]].<ref name=Bond2012>{{cite book |first=W. J. |last=Bond |chapter=11. Keystone species |editor-last1=Schulze |editor-first1=Ernst-Detlef |editor-last2=Mooney |editor-first2=Harold A. |title=Biodiversity and Ecosystem Function |date=2012 |publisher=Springer |isbn=978-3-642-58001-7 |page=237}}</ref> Introduction or removal of this predator, or changes in its population density, can have drastic cascading effects on the equilibrium of many other populations in the ecosystem. For example, grazers of a grassland may prevent a single dominant species from taking over.<ref>{{cite book |author1=Botkin, D. |author2=Keller, E. |date=2003 |title=Environmental Science: Earth as a living planet |publisher=John Wiley & Sons |isbn=978-0-471-38914-9 |page=2}}</ref>

[[File:Fig. 1 -Riparian willow recovery (26485120926) horiz.jpg|thumb|center|upright=2.8<!--double image-->|Riparian willow recovery at Blacktail Creek, [[Yellowstone National Park]], after reintroduction of wolves, the local [[keystone species]] and [[apex predator]].<ref name="Ripple2004"/> Left, in 2002; right, in 2015]]

The elimination of wolves from [[Yellowstone National Park]] had profound impacts on the [[trophic pyramid]]. In that area, wolves are both keystone species and apex predators. Without predation, herbivores began to over-graze many woody browse species, affecting the area's plant populations. In addition, wolves often kept animals from grazing near streams, protecting the [[beaver]]s' food sources. The removal of wolves had a direct effect on the beaver population, as their habitat became territory for grazing. Increased browsing on [[willow]]s and [[conifer]]s along Blacktail Creek due to a lack of predation caused channel incision because the reduced beaver population was no longer able to slow the water down and keep the soil in place. The predators were thus demonstrated to be of vital importance in the ecosystem.<ref name="Ripple2004">{{cite journal |last1=Ripple |first1=William J. |last2=Beschta |first2=Robert L. |title=Wolves and the Ecology of Fear: Can Predation Risk Structure Ecosystems? |journal=BioScience |volume=54 |issue=8 |year=2004 |doi=10.1641/0006-3568(2004)054[0755:WATEOF]2.0.CO;2 | page=755|doi-access=free }}</ref>

=== Population dynamics ===
{{further|Population dynamics|Lotka–Volterra equations}}

[[File:Milliers fourrures vendues en environ 90 ans odum 1953 en.jpg|right|upright=1.5|thumb|Numbers of [[snowshoe hare]] (''Lepus americanus'') (yellow background) and Canada lynx (black line, foreground) furs sold to the [[Hudson's Bay Company]] from 1845 to 1935|alt=A line graph of the number of Canada lynx furs sold to the Hudson's Bay Company on the vertical axis against the numbers of snowshoe hare on the horizontal axis for the period 1845 to 1935]]

In the absence of predators, the population of a species can grow exponentially until it approaches the [[carrying capacity]] of the environment.<ref>{{cite book |last1=Neal |first1=Dick |title=Introduction to population biology |date=2004 |publisher=Cambridge University Press |isbn=978-0-521-53223-5 |pages=68–69}}</ref> Predators limit the growth of prey both by consuming them and by changing their behavior.<ref>{{cite journal |last1=Nelson |first1=Erik H. |last2=Matthews |first2=Christopher E. |last3=Rosenheim |first3=Jay A. |title=Predators Reduce Prey Population Growth by Inducing Changes in Prey Behavior |journal=Ecology |date=July 2004 |volume=85 |issue=7 |pages=1853–1858 |jstor=3450359 |doi=10.1890/03-3109 |bibcode=2004Ecol...85.1853N |url=https://rosenheim.faculty.ucdavis.edu/wp-content/uploads/sites/137/2014/09/Ecology2004ms.pdf }}</ref> Increases or decreases in the prey population can also lead to increases or decreases in the number of predators, for example, through an increase in the number of young they bear.

Cyclical fluctuations have been seen in populations of predator and prey, often with offsets between the predator and prey cycles. A well-known example is that of the [[snowshoe hare]] and [[lynx]]. Over a broad span of [[boreal forest]]s in Alaska and Canada, the hare populations fluctuate in near synchrony with a 10-year period, and the lynx populations fluctuate in response. This was first seen in historical records of animals caught by [[Fur trade|fur hunter]]s for the [[Hudson's Bay Company]] over more than a century.<ref name=Krebs>{{cite journal |last1=Krebs |first1=Charles J. |last2=Boonstra |first2=Rudy |last3=Boutin |first3=Stan |last4=Sinclair |first4=A.R.E. |title=What Drives the 10-year Cycle of Snowshoe Hares? |journal=BioScience |date=2001 |volume=51 |issue=1 |page=25 |doi=10.1641/0006-3568(2001)051[0025:WDTYCO]2.0.CO;2|doi-access=free |hdl=1807/359 |hdl-access=free }}</ref><ref name=classics>{{cite journal |last1=Peckarsky |first1=Barbara L. |last2=Abrams |first2=Peter A. |last3=Bolnick |first3=Daniel I. |last4=Dill |first4=Lawrence M. |last5=Grabowski |first5=Jonathan H. |last6=Luttbeg |first6=Barney |last7=Orrock |first7=John L. |last8=Peacor |first8=Scott D. |last9=Preisser |first9=Evan L. |last10=Schmitz |first10=Oswald J. |last11=Trussell |first11=Geoffrey C. |title=Revisiting the classics: considering nonconsumptive effects in textbook examples of predator–prey interactions |journal=Ecology |date=September 2008 |volume=89 |issue=9 |pages=2416–2425 |doi=10.1890/07-1131.1 |pmid=18831163 |bibcode=2008Ecol...89.2416P |url=https://digitalcommons.uri.edu/bio_facpubs/121 }}</ref><ref name="The Snowshoe Hare 10-year Cycle – A Cautionary Tale">{{cite web |last1=Krebs |first1=Charley |last2=Myers |first2=Judy |title=The Snowshoe Hare 10-year Cycle – A Cautionary Tale |url=https://www.zoology.ubc.ca/~krebs/ecological_rants/?p=786 |website=Ecological rants |publisher=University of British Columbia |access-date=2 October 2018|date=12 July 2014 }}</ref><ref name="BBC Bitesize - Predators and their prey">{{cite web |title=Predators and their prey |url=https://www.bbc.co.uk/schools/gcsebitesize/science/ocr_gateway/understanding_environment/interdependencerev2.shtml |archive-url=https://web.archive.org/web/20120502225541/http://www.bbc.co.uk/schools/gcsebitesize/science/ocr_gateway/understanding_environment/interdependencerev2.shtml |archive-date=2 May 2012 |website=BBC Bitesize |publisher=[[BBC]] |access-date=7 October 2015}}</ref>

[[File:Lotka-Volterra model (1.1, 0.4, 0.4, 0.1).png|thumb|upright=1.35|left|[[Predator]]-prey population cycles in a [[Lotka–Volterra equations|Lotka–Volterra model]]<!-- with the parameters set to (1.1, 0.4, 0.4, 0.1)-->]]

A simple model of a system with one species each of predator and prey, the [[Lotka–Volterra equations]], predicts population cycles.<ref name="Goel1971">{{cite book |last1=Goel |first1=Narendra S. |first2=S. C. |last2=Maitra |first3=E. W. |last3=Montroll |title=On the Volterra and Other Non-Linear Models of Interacting Populations |publisher=Academic Press |year=1971 |isbn=978-0-12-287450-5}}</ref> However, attempts to reproduce the predictions of this model in the laboratory have often failed; for example, when the protozoan ''[[Didinium#Didinium nasutum|Didinium nasutum]]'' is added to a culture containing its prey, ''[[Paramecium caudatum]]'', the latter is often driven to extinction.<ref name=Princeton>{{cite book |last1=Levin |first1=Simon A. |last2=Carpenter |first2=Stephen R. |last3=Godfray |first3=H. Charles J. |last4=Kinzig |first4=Ann P. |last5=Loreau |first5=Michel |last6=Losos |first6=Jonathan B. |last7=Walker |first7=Brian |last8=Wilcove |first8=David S. |title=The Princeton guide to ecology |url=https://archive.org/details/princetonguideto00levi |url-access=limited |date=2009 |publisher=Princeton University Press |isbn=978-1-4008-3302-3 |pages=[https://archive.org/details/princetonguideto00levi/page/n218 204]–209}}</ref>

The Lotka–Volterra equations rely on several simplifying assumptions, and they are [[structural stability|structurally unstable]], meaning that any change in the equations can stabilize or destabilize the dynamics.<ref name=Murdoch>{{cite book |last1=Murdoch |first1=William W. |last2=Briggs |first2=Cheryl J. |last3=Nisbet |first3=Roger M. |title=Consumer-resource dynamics |date=2013 |publisher=Princeton University Press |isbn=978-1-4008-4725-9|page=39}}</ref><ref>{{cite book |last1=Nowak |first1=Martin |last2=May |first2=Robert M. |title=Virus Dynamics: Mathematical Principles of Immunology and Virology |date=2000 |publisher=Oxford University Press |isbn=978-0-19-158851-8 |page=8}}</ref> For example, one assumption is that predators have a linear [[functional response]] to prey: the rate of kills increases in proportion to the rate of encounters. If this rate is limited by time spent handling each catch, then prey populations can reach densities above which predators cannot control them.<ref name=Princeton/> Another assumption is that all prey individuals are identical. In reality, predators tend to select young, weak, and ill individuals, leaving prey populations able to regrow.<ref name="Genovart2010">{{cite journal |last1=Genovart |first1=M. |last2=Negre |first2=N. |last3=Tavecchia |first3=G. |last4=Bistuer |first4=A. |last5=Parpal |first5=L. |last6=Oro |first6=D. |year=2010 |title=The young, the weak and the sick: evidence of natural selection by predation |journal=PLOS ONE|volume=5 |issue=3 | article-number=e9774 |doi=10.1371/journal.pone.0009774 |pmid=20333305 |pmc=2841644|bibcode=2010PLoSO...5.9774G |doi-access=free }}</ref>

Many factors can stabilize predator and prey populations.<ref>{{harvnb|Rockwood|2009|page=281}}</ref> One example is the presence of multiple predators, particularly generalists that are attracted to a given prey species if it is abundant and look elsewhere if it is not.<ref>{{harvnb|Rockwood|2009|page=246}}</ref> As a result, population cycles tend to be found in northern temperate and [[subarctic]] ecosystems because the food webs are simpler.<ref>{{harvnb|Rockwood|2009|pages=271–272}}</ref> The snowshoe hare-lynx system is subarctic, but even this involves other predators, including coyotes, [[goshawk]]s and [[great horned owl]]s, and the cycle is reinforced by variations in the food available to the hares.<ref>{{harvnb|Rockwood|2009|page=272–273}}</ref>

A range of mathematical models have been developed by relaxing the assumptions made in the Lotka–Volterra model; these variously allow animals to have [[spatial distribution|geographic distribution]]s, or to [[animal migration|migrate]]; to have differences between individuals, such as [[sex]]es and an [[age structure]], so that only some individuals reproduce; to live in a varying environment, such as with changing [[season]]s;<ref name="Cushing2005">{{cite journal |last1=Cushing |first1=J. M. |title=Book Review: Mathematics in population biology |journal=Bulletin of the American Mathematical Society |date=30 March 2005 |volume=42 |issue=4 |pages=501–506 |doi=10.1090/S0273-0979-05-01055-4 |doi-access=free }}</ref><ref>{{cite book|last=Thieme|first=Horst R. |title=Mathematics in Population Biology|url=https://books.google.com/books?id=cHcjnkrMweYC |year=2003 |publisher=Princeton University Press |isbn=978-0-691-09291-1}}</ref> and analysing the interactions of more than just two species at once. Such models predict widely differing and often [[chaos theory|chaotic]] predator-prey population dynamics.<ref name="Cushing2005"/><ref name="KozlovVakulenko2013">{{cite journal |last1=Kozlov |first1=Vladimir |last2=Vakulenko |first2=Sergey |title=On chaos in Lotka–Volterra systems: an analytical approach |journal=Nonlinearity |volume=26 |issue=8 | date=3 July 2013 |doi=10.1088/0951-7715/26/8/2299 |pages=2299–2314|bibcode=2013Nonli..26.2299K | s2cid=121559550 }}</ref> The presence of [[refuge (ecology)|refuge areas]], where prey are safe from predators, may enable prey to maintain larger populations but may also destabilize the dynamics.<ref>{{cite journal |doi=10.1016/0040-5809(87)90019-0 |title=Prey refuges and predator-prey stability |journal=Theoretical Population Biology |volume=31 |pages=1–12 |year=1987 |last1=Sih |first1=Andrew |issue=1 |bibcode=1987TPBio..31....1S }}</ref><ref>{{cite journal |doi=10.1016/0040-5809(86)90004-3 |pmid=3961711 |title=The effects of refuges on predator-prey interactions: A reconsideration |journal=Theoretical Population Biology |volume=29 |issue=1 |pages=38–63 |year=1986 |last1=McNair |first1=James N |bibcode=1986TPBio..29...38M }}</ref><ref name="BerrymanHawkins2006">{{cite journal |last1=Berryman |first1=Alan A. |last2=Hawkins |first2=Bradford A. |last3=Hawkins |first3=Bradford A. |title=The refuge as an integrating concept in ecology and evolution |journal=Oikos |volume=115 |issue=1 |year=2006 |pages=192–196 |doi=10.1111/j.0030-1299.2006.15188.x|bibcode=2006Oikos.115..192B }}</ref><ref>{{cite journal |doi=10.1016/j.tpb.2009.08.005 |pmid=19751753 |title=A predator–prey refuge system: Evolutionary stability in ecological systems |journal=Theoretical Population Biology |volume=76 |issue=4 |pages=248–57 |year=2009 |last1=Cressman |first1=Ross |last2=Garay |first2=József |bibcode=2009TPBio..76..248C }}</ref>

== Evolutionary history ==

{{further|History of life}}

Predation dates from before the rise of commonly recognized carnivores by hundreds of millions (perhaps billions) of years. Predation has evolved repeatedly in different groups of organisms.<ref name="Bengtson2002"/><ref>{{cite journal |author=Abrams, P. A. |date=2000 |title=The evolution of predator-prey interactions: theory and evidence |journal=Annual Review of Ecology and Systematics |volume=31 |issue=1 |pages=79–105|doi=10.1146/annurev.ecolsys.31.1.79 |bibcode=2000AnRES..31...79A }}</ref> The rise of [[eukaryotic]] cells at around 2.7 Gya, the rise of multicellular organisms at about 2 Gya, and the rise of mobile predators (around 600 Mya - 2 Gya, probably around 1 Gya) have all been attributed to early predatory behavior, and many very early remains show evidence of boreholes or other markings attributed to small predator species.<ref name=Bengtson2002 /> It likely triggered major evolutionary transitions including the arrival of [[Cell (biology)|cell]]s, [[eukaryote]]s, [[sexual reproduction]], [[multicellularity]], increased size, mobility (including [[insect flight]]<ref name=GrimaldiEngel2005/>) and armoured shells and exoskeletons.<ref name="Bengtson2002"/>

The earliest predators were microbial organisms, which engulfed or grazed on others. Because the fossil record is poor, these first predators could date back anywhere between 1 and over 2.7 Gya (billion years ago).<ref name=Bengtson2002/> Predation visibly became important shortly before the [[Cambrian]] period—around {{Ma |550}}—as evidenced by the almost simultaneous development of [[calcification]] in animals and algae,<ref name="Grant1991">{{cite journal |author1=Grant, S. W. F. |author2=Knoll, A. H. |author3=Germs, G. J. B. |year=1991 |title=Probable Calcified Metaphytes in the Latest Proterozoic Nama Group, Namibia: Origin, Diagenesis, and Implications |journal=Journal of Paleontology |volume=65 |issue=1 |pages=1–18 |pmid=11538648 |jstor=1305691|doi=10.1017/S002233600002014X |bibcode=1991JPal...65....1G |s2cid=26792772 }}</ref> and predation-avoiding [[burrow]]ing. However, predators had been grazing on micro-organisms since at least {{Ma |1000}},<ref name="Bengtson2002">{{cite book |author=Bengtson, S. |year=2002 |contribution=Origins and early evolution of predation |title=The fossil record of predation. The Paleontological Society Papers 8 |editor-last1=Kowalewski |editor-first1=M. |editor-last2=Kelley |editor-first2=P. H. |pages=289–317 |publisher=The Paleontological Society |url=http://www.nrm.se/download/18.4e32c81078a8d9249800021552/Bengtson2002predation.pdf |access-date=23 July 2008 |archive-date=10 September 2008 |archive-url=https://web.archive.org/web/20080910205539/http://www.nrm.se/download/18.4e32c81078a8d9249800021552/Bengtson2002predation.pdf }}</ref><ref name="Awramik1971">{{cite journal |author=Awramik, S. M. |title=Precambrian columnar stromatolite diversity: Reflection of metazoan appearance |journal=Science |volume=174  |issue=4011 |pages=825–827 |date=19 November 1971 |doi=10.1126/science.174.4011.825 |pmid=17759393 |bibcode=1971Sci...174..825A|s2cid=2302113 }}</ref><ref name="Stanley2008">{{cite journal |title=Predation defeats competition on the seafloor |author=Stanley, Steven M. |year=2008 |journal=Paleobiology |volume=34 |pages=1–21 |doi=10.1666/07026.1 |issue=1|bibcode=2008Pbio...34....1S |s2cid=83713101 }}</ref> with evidence of selective (rather than random) predation from a similar time.<ref>{{cite journal |doi=10.1042/ETLS20170153 |title=Implications of selective predation on the macroevolution of eukaryotes: Evidence from Arctic Canada |journal=Emerging Topics in Life Sciences|volume=2 |issue=2 |pages=247–255 |year=2018 |last1=Loron |first1=Corentin C. |last2=Rainbird |first2=Robert H. |last3=Turner |first3=Elizabeth C. |last4=Wilder Greenman |first4=J. |last5=Javaux |first5=Emmanuelle J.|pmid=32412621 |s2cid=92505644 }}</ref>

''[[Auroralumina attenboroughii]]'' is an Ediacaran crown-group [[cnidaria]]n (557–562 mya, some 20 million years before the Cambrian explosion) from [[Charnwood Forest]], England. It is thought to be one of the earliest predatory animals, catching small prey with its [[nematocyst]]s as modern cnidarians do.<ref name="Dunn Kenchington Parry Clark 2022">{{cite journal |last1=Dunn |first1=F. S. |last2=Kenchington |first2=C. G. |last3=Parry |first3=L. A. |last4=Clark |first4=J. W. |last5=Kendall |first5=R. S. |last6=Wilby |first6=P. R. |title=A crown-group cnidarian from the Ediacaran of Charnwood Forest, UK |journal=Nature Ecology & Evolution |date=25 July 2022 |volume=6 |issue=8 |pages=1095–1104 |doi=10.1038/s41559-022-01807-x |pmid=35879540 |pmc=9349040 |bibcode=2022NatEE...6.1095D }}</ref>

The [[fossil record]] demonstrates a long history of interactions between predators and their prey from the Cambrian period onwards, showing for example that some predators drilled through the shells of [[bivalve]] and [[gastropod]] molluscs, while others ate these organisms by breaking their shells.<ref name="Kelley">{{cite book |last=Kelley |first=Patricia |title=Predator—Prey Interactions in the Fossil Record |publisher=Springer |year=2003 | isbn=978-1-4615-0161-9 | oclc=840283264 |pages=113–139, 141–176 and passim}}</ref>
Among the Cambrian predators were invertebrates like the [[anomalocaridid]]s with appendages suitable for grabbing prey, large compound eyes and jaws made of a hard material like that in the [[exoskeleton]] of an insect.<ref>{{cite journal |last=Daley |first=Allison C. |title=Anomalocaridids |journal=Current Biology |volume=23 |issue=19 |pages=R860–R861 |doi=10.1016/j.cub.2013.07.008|pmid=24112975 |year=2013 |doi-access=free |bibcode=2013CBio...23.R860D }}</ref>
Some of the first [[jawed fish|fish to have jaws]] were the armoured and mainly predatory [[placoderms]] of the [[Silurian]] to [[Devonian]] periods, one of which, the {{convert|6|m|abbr=on}} ''[[Dunkleosteus]]'', is considered the world's first [[vertebrate]] "superpredator", preying upon other predators.<ref name=Anderson&Westneat2009>{{cite journal |last=Anderson |first=P. S. L. |author2=Westneat, M. |title=A biomechanical model of feeding kinematics for Dunkleosteus terrelli (Arthrodira, Placodermi) |journal=Paleobiology |date=2009 |volume=35 |issue=2 |pages=251–269|doi=10.1666/08011.1|bibcode=2009Pbio...35..251A |s2cid=86203770 }}</ref><ref name="Carr2010">{{cite journal |last1=Carr |first1=Robert K. |title=Paleoecology of Dunkleosteus Terrelli (Placodermi: Arthrodira) |journal=Kirtlandia |date=2010 |volume=57 |pages=36–45 |url=https://biostor.org/reference/193162 }}</ref>
[[Insect]]s developed the ability to fly in the Early [[Carboniferous]] or Late Devonian, enabling them among other things to escape from predators.<ref name=GrimaldiEngel2005>{{cite book |last1=Grimaldi |first1=David |author-link1=David Grimaldi (entomologist) |last2=Engel |first2=Michael S. |author-link2=Michael S. Engel |year=2005 |title=Evolution of the Insects |url=https://archive.org/details/evolutioninsects00grim_110 |url-access=limited |publisher=Cambridge University Press |isbn=978-0-521-82149-0 |pages=[https://archive.org/details/evolutioninsects00grim_110/page/n169 155]–160}}</ref>
Among the largest predators that have ever lived were the [[theropod dinosaur]]s such as ''[[Tyrannosaurus]]'' from the [[Cretaceous]] period. They preyed upon herbivorous dinosaurs such as [[hadrosaur]]s, [[ceratopsia]]ns and [[ankylosaur]]s.<ref>{{cite journal |last1=Sampson |first1=Scott D. |last2=Loewen |first2=Mark A. |title=Tyrannosaurus rex from the Upper Cretaceous (Maastrichtian) North Horn Formation of Utah: biogeographic and paleoecologic implications |journal=Journal of Vertebrate Paleontology |date=27 June 2005 |volume=25 |issue=2 |pages=469–472 |doi=10.1671/0272-4634(2005)025[0469:TRFTUC]2.0.CO;2 |s2cid=131583311 }}</ref>

<gallery class="center" mode="nolines" widths="210px" heights="180px">
File:Auroralumina attenboroughii reconstruction.jpg|''[[Auroralumina attenboroughii]]'', an [[Ediacaran]] predator (c. 560 mya). It was a stem-group [[cnidarian]], catching prey with its [[nematocyst]]s.<ref name="Dunn Kenchington Parry Clark 2022"/>
File:Cambrian substrate revolution 02.png|The [[Cambrian substrate revolution]] saw life on the [[sea floor]] change from minimal burrowing (left) to a diverse burrowing fauna (right), probably to avoid new [[Cambrian]] predators.
File:20191021 Peytoia nathorsti Laggania cambria.png|The [[anomalocaridid]] ''[[Peytoia]]'', a [[Cambrian]] invertebrate, probably an apex predator
File:Dunkleosteus terrelli (2024).png|[[Dunkleosteus|Dunkleosteus terrelli]], a [[Devonian]] [[placoderm]], perhaps the world's first vertebrate [[Super-predator|superpredator]]
File:Meganeura.png|''[[Meganeura monyi]]'', a predatory [[Carboniferous]] [[insect]] related to [[Dragonfly|dragonflies]], could fly to escape terrestrial predators. Its large size, with a wingspan of {{convert|65|cm|in|-1|abbr=on}}, may reflect the lack of vertebrate aerial predators at that time.
File:Tyrannosaurus rex Reconstruction by Nobu Tamura.jpg|''[[Tyrannosaurus]]'', a large [[theropod dinosaur]] of the [[Cretaceous]]
</gallery>

==In human society==

{{further|Human uses of animals}}

[[File:Bushmen hunters.jpg|thumb|upright|[[San people|San]] hunter, Botswana]]

===Practical uses===

Humans, as [[omnivore]]s, are to some extent predatory,<ref name="DarimontFoxBryanReimchen2015">{{cite journal |last1=Darimont |first1=C. T. |last2=Fox |first2=C. H. |last3=Bryan |first3=H. M. |last4=Reimchen |first4=T. E. |title=The unique ecology of human predators |journal=Science |volume=349 |issue=6250 | date=20 August 2015 |doi=10.1126/science.aac4249 |pmid=26293961 |pages=858–860 |bibcode=2015Sci...349..858D| s2cid=4985359 }}</ref> using weapons and tools to [[fishing|fish]],<ref>{{cite book|first=Otto |last=Gabriel |author2=von Brandt, Andres |year=2005 |publisher=Blackwell|title=Fish catching methods of the world | isbn=978-0-85238-280-6}}</ref> [[hunting|hunt]] and [[animal trapping|trap]] animals.<ref>{{cite book|last1=Griffin |first1=Emma |title=Blood Sport: Hunting in Britain Since 1066 |date=2008 |publisher=Yale University Press|isbn=978-0-300-14545-8}}</ref> They also use other predatory species such as [[Hunting dog|dogs]], [[cormorant]]s,<ref>{{cite book |author=King, Richard J. |title=The Devil's Cormorant: A Natural History |url=https://books.google.com/books?id=ucGyAAAAQBAJ&pg=PA9 |date=1 October 2013 |publisher=University of New Hampshire Press |isbn=978-1-61168-225-0 |page=9}}</ref> and [[falcon]]s to catch prey for food or for sport.<ref>{{cite book |last1=Glasier |first1=Phillip |title=Falconry and Hawking |date=1998 |publisher=Batsford |isbn=978-0-7134-8407-6}}</ref> 
Two mid-sized predators, dogs and cats, are the animals most often kept as [[pet]]s in western societies.<ref name="Aegerter Fouracre Smith p=e0174709">{{cite journal |last1=Aegerter |first1=James |last2=Fouracre |first2=David |last3=Smith |first3=Graham C. | editor-last=Olsson | editor-first=I Anna S |title=A first estimate of the structure and density of the populations of pet cats and dogs across Great Britain |journal=PLOS ONE|volume=12 |issue=4 | date=2017 |doi=10.1371/journal.pone.0174709 |pmid=28403172 |pmc=5389805 | article-number=e0174709|bibcode=2017PLoSO..1274709A |doi-access=free }}</ref><ref>{{cite web |last=The Humane Society of the United States |title=U.S. Pet Ownership Statistics |url=http://www.humanesociety.org/issues/pet_overpopulation/facts/pet_ownership_statistics.html |access-date=27 April 2012}}</ref>
Human hunters, including the [[San people|San]] of southern Africa, use [[persistence hunting]], a form of pursuit predation where the pursuer may be slower than prey such as a [[kudu]] antelope over short distances, but follows it in the midday heat until it is exhausted, a pursuit that can take up to five hours.<ref name="Liebenberg20089">{{cite journal |last=Liebenberg |first=Louis |title=The relevance of persistence hunting to human evolution |journal=Journal of Human Evolution |year=2008 |volume=55 |issue=6 |pages=1156–1159 |doi=10.1016/j.jhevol.2008.07.004 |pmid=18760825 |bibcode=2008JHumE..55.1156L }}</ref><ref>{{cite web |url=https://www.bbc.co.uk/pressoffice/pressreleases/stories/2002/10_october/31/progs6-10_mammalspack.pdf |title=Food For Thought |publisher=[[British Broadcasting Corporation]] |work=[[The Life of Mammals]] |date=31 October 2002}}</ref> 
 
In [[biological pest control]], predators (and parasitoids) from a pest's natural range are introduced to control populations, at the risk of causing unforeseen problems. Natural predators, provided they do no harm to non-pest species, are an environmentally friendly and sustainable way of reducing damage to crops and an alternative to the use of chemical agents such as [[pesticide]]s.<ref>{{cite book |last1=Flint |first1=Mary Louise |last2=Dreistadt |first2=Steve H. |title=Natural Enemies Handbook: The Illustrated Guide to Biological Pest Control |date=1998 |publisher=University of California Press |isbn=978-0-520-21801-7 }}{{pn|date=July 2024}}</ref>

===Symbolic uses===

[[File:Lupa Capitolina, Rome.jpg|thumb|The [[Capitoline Wolf]] suckling [[Romulus and Remus]], the mythical founders of [[Ancient Rome|Rome]]]]

In film, the idea of the predator as a dangerous if [[humanoid]] enemy is used in the 1987 [[science fiction]] [[horror film|horror]] [[action film]] [[Predator (film)|''Predator'']] and [[Predator (franchise)|its three sequels]].<ref>{{cite book |last=Johnston |first=Keith M. |year=2013 |title=Science Fiction Film: A Critical Introduction |publisher=[[Berg Publishers]] |isbn=978-0-85785-056-0 |page=98 |url=https://books.google.com/books?id=yUAfAAAAQBAJ&pg=PA98}}</ref><ref>{{cite magazine |last1=Newby |first1=Richard |title=Is 'Predator' Finally Getting a Worthy Sequel? |url=https://www.hollywoodreporter.com/heat-vision/predator-is-poised-be-franchises-first-good-sequel-1111354 |magazine=Hollywood Reporter |access-date=7 September 2018 |date=13 May 2018}}</ref> A terrifying predator, a gigantic [[Man-eating animal|man-eating]] [[great white shark]], is central, too, to [[Steven Spielberg]]'s 1974 thriller ''[[Jaws (film)|Jaws]]''.<ref>{{cite book |last=Schatz |first=Thomas |chapter=The New Hollywood |title=Movie Blockbusters |page=[https://books.google.com/books?id=6Q-OAQAAQBAJ&pg=PA25&dq=%22jaws+was+essentially+an+action+film+and+a+thriller%22 25]}} In: {{cite book |last=Stringer |first=Julian |title=Movie Blockbusters |isbn=978-0-415-25608-7 |publisher=[[Routledge]] |year=2003 |pages=15–44}}</ref>

Among poetry on the theme of predation, a predator's consciousness might be explored, such as in [[Ted Hughes]]'s ''Pike''.<ref>{{cite news |last1=Davison |first1=Peter |title=Predators and Prey {{!}} Selected Poems, 1957–1994 by Ted Hughes |url=https://www.nytimes.com/2002/12/01/books/predators-and-prey.html |access-date=5 October 2018 |work=[[The New York Times]] |date=1 December 2002 |quote=Hughes's earliest books contained a bewildering profusion of poems between their covers: ... fish and fowl, beasts of the field and forest, vigorous embodiments of predators and prey. Hughes as a student had taken up anthropology, not literature, and he chose to meditate his way into trancelike states of preconsciousness before committing poems to paper. His poems, early or late, enter into the relations of living creatures; they move in close to animal consciousness: ''The Thought-Fox,'' ''Esther's Tomcat,'' ''Pike.''}}</ref> The phrase "Nature, red in tooth and claw" from [[Alfred, Lord Tennyson]]'s 1849 poem "[[In Memoriam A.H.H.]]" has been interpreted as referring to the struggle between predators and prey.<ref name="Gould 1995">{{cite book |last=Gould |first=Stephen Jay |author-link=Stephen Jay Gould |chapter =The Tooth and Claw Centennial |title =Dinosaur in a Haystack |date=1995 |publisher=Harmony Books |pages=63–75 |isbn=978-0-517-70393-9}}</ref>

In mythology and folk fable, predators such as the fox and wolf have mixed reputations.<ref name=Wallner1998>{{cite web |last1=Wallner |first1=Astrid |title=The role of predators in Mythology |url=https://www.waldwissen.net/wald/tiere/saeuger/wsl_raubtiere_mythologie/index_EN |publisher=WaldWissen Information for Forest Management |access-date=5 October 2018 |date=18 July 2005 |archive-date=5 October 2018 |archive-url=https://web.archive.org/web/20181005154029/https://www.waldwissen.net/wald/tiere/saeuger/wsl_raubtiere_mythologie/index_EN }} translated from Wallner, A. (1998) ''Die Bedeutung der Raubtiere in der Mythologie: Ergebnisse einer Literaturstudie''. – Inf.bl. Forsch.bereiches Landsch.ökol. 39: 4–5.</ref> The fox was a symbol of fertility in ancient Greece, but a weather demon in northern Europe, and a creature of the devil in early Christianity; the fox is presented as sly, greedy, and cunning in fables from [[Aesop's Fables|Aesop]] onwards.<ref name=Wallner1998/> The big bad wolf is known to children in tales such as ''[[Little Red Riding Hood]]'', but is a demonic figure in the Icelandic [[Edda]] sagas, where the wolf [[Fenrir]] appears in the apocalyptic [[Ragnarok|ending of the world]].<ref name=Wallner1998/> In the Middle Ages, belief spread in [[werewolf|werewolves]], men transformed into wolves.<ref name=Wallner1998/> In ancient Rome, and in ancient Egypt, the wolf was worshipped, the she-wolf appearing in the founding myth of Rome, suckling [[Romulus and Remus]].<ref name=Wallner1998/> More recently, in [[Rudyard Kipling]]'s 1894 ''[[The Jungle Book]]'', Mowgli is raised by the wolf pack.<ref name=Wallner1998/> Attitudes to large predators in North America, such as wolf, [[grizzly bear]] and cougar, have shifted from hostility or ambivalence, accompanied by active persecution, towards positive and protective in the second half of the 20th century.<ref name="KellertBlackRushBath1996">{{cite journal |last1=Kellert |first1=Stephen R. |last2=Black |first2=Matthew |last3=Rush |first3=Colleen Reid |last4=Bath |first4=Alistair J. |title=Human Culture and Large Carnivore Conservation in North America |journal=Conservation Biology |volume=10 |issue=4 |year=1996 |doi=10.1046/j.1523-1739.1996.10040977.x |pages=977–990|bibcode=1996ConBi..10..977K }}
</ref>

== See also ==

* [[Ecology of fear (concept)|Ecology of fear]]
* [[Predation problem]]
* [[Predator–prey reversal]]
* [[Prey naiveté]]
* [[Wa-Tor]]
* [[Cannibalism]]

== Notes ==

{{notelist}}

== References ==

{{reflist}}

== Sources ==

* {{cite book |last1=Beauchamp |first1=Guy |title=Social predation: how group living benefits predators and prey |date=2012 |publisher=Elsevier |isbn=978-0-12-407654-9 }}
* {{cite book |last1=Bell |first1=W. J. |title=Searching Behaviour: the behavioural ecology of finding resources |date=2012 |publisher=Springer Netherlands |isbn=978-94-011-3098-1}}
* {{cite book |last=Caro |first=Tim |author-link=Tim Caro |title=Antipredator Defenses in Birds and Mammals |date=2005 |publisher=University of Chicago Press |isbn=978-0-226-09436-6}}
* {{cite book |last=Cott |first=Hugh B. |author-link=Hugh B. Cott |year=1940 |title=Adaptive Coloration in Animals |publisher=Methuen |title-link=Adaptive Coloration in Animals |oclc=974070031 }}
* {{cite book |last1=Jacobs |first1=David Steve |last2=Bastian |first2=Anna |title=Predator-prey interactions: co-evolution between bats and their prey |date=2017 |publisher=Springer |isbn=978-3-319-32492-0}}
* {{cite book |last1=Rockwood |first1=Larry L. |title=Introduction to population ecology |date=2009 |publisher=John Wiley & Sons |isbn=978-1-4443-0910-2|page=281}}
* {{cite book |last1=Ruxton |first1=Graeme D. |author1-link=Graeme Ruxton |last2=Sherratt |first2=Tom N. |author2-link=Thomas N. Sherratt |last3=Speed |first3=Michael P. |title=Avoiding attack: the evolutionary ecology of crypsis, warning signals, and mimicry |date=2004 |publisher=Oxford University Press |isbn=978-0-19-852859-3 }}

== External links ==
* {{Wikiquote-inline}}
* {{Commons category-inline|Predation}}

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[[Category:Ecology]]
[[Category:Predation| ]]
[[Category:Biological pest control]]