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Predation is a
biological interaction In ecology Ecology (from el, οἶκος, "house" and el, -λογία, label=none, "study of") is the study of the relationships between living organisms, including humans, and their physical environment. Topics of interest include the bio ...
where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common
feeding behaviours Feeding is the process by which organisms, typically animals, obtain food. Terminology often uses either the suffixes -vore, -vory, or -vorous from Latin ''vorare'', meaning "to devour", or -phage, -phagy, or -phagous from Greek φαγε ...
that includes
parasitism Parasitism is a close relationship between species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, ...

parasitism
and
micropredation Parasitism is a symbiotic Symbiosis (from Ancient Greek, Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different Organism, biological or ...
(which usually do not kill the
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places *Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman *Michel Host (19 ...
) and
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable stra ...
ism (which always does, eventually). It is distinct from
scavenging Scavengers are animals that consume dead organisms that have died from causes other than predation. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding behavior. Scavengers play an important role ...
on dead prey, though many predators also scavenge; it overlaps with
herbivory File:Land_Snail_radula_tracks.jpg#, 250px, Tracks made by terrestrial gastropods with their radulas, scraping green algae from a surface inside a greenhouse A herbivore is an animal anatomically and physiologically adapted to eating plant mater ...
, as
seed predator A aggregate accessory fruit damaged by a mouse eating the seeds (achenes). Seed predation, often referred to as granivory, is a type of plant-animal interaction in which granivores (seed predators) feed on the seeds of plants as a main or exclu ...
s and destructive
frugivore A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance and ...
s are predators. Predators may actively search for or pursue prey or wait for it, often concealed. When prey is detected, the predator assesses whether to attack it. This may involve
ambush An ambush is a long-established military tactics, military tactic in which combatants take advantage of concealment or the element of surprise to attack unsuspecting enemy combatants from concealed positions, such as among dense underbrush o ...
or
pursuit predation Pursuit predation is a form of predation Predation is a biological interaction In ecology Ecology (from el, οἶκος, "house" and el, -λογία, label=none, "study of") is the study of the relationships between living organisms ...
, sometimes after stalking the prey. If the attack is successful, the predator kills the prey, removes any inedible parts like the shell or spines, and eats it. Predators are adapted and often highly specialized for hunting, with acute senses such as
vision Vision or The Vision may refer to: Perception Optical perception * Visual perception Visual perception is the ability to interpret the surrounding environment (biophysical), environment through photopic vision (daytime vision), color visio ...

vision
,
hearing Schematic diagram of the human ear Hearing, or auditory perception, is the ability to perceive Sound, sounds through an organ, such as an ear, by detecting Vibration, vibrations as periodic changes in the pressure of a surrounding medium. Th ...

hearing
, or smell. Many predatory
animal Animals (also called Metazoa) are multicellular A multicellular organism is an organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are composed of cells ...

animal
s, both
vertebrate Vertebrates () comprise all species of animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic ma ...
and
invertebrate Invertebrates are animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This includes all animals apart from the chordata, chordate subphylum vertebrate, Vertebra ...
, have sharp
claw A domestic cat's retractable claw in protracted position A claw is a curved, pointed appendage found at the end of a toe or finger in most amniote Amniotes (from Greek ἀμνίον ''amnion'', "membrane surrounding the fetus", earlier "bo ...

claw
s or
jaw The jaw is any opposable articulated structure at the entrance of the , typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serving t ...

jaw
s to grip, kill, and cut up their prey. Other adaptations include stealth and
aggressive mimicry Aggressive mimicry is a form of mimicry In evolutionary biology Evolutionary biology is the subfield of biology that studies the evolution, evolutionary processes (natural selection, common descent, speciation) that produced the Biod ...
that improve hunting efficiency. Predation has a powerful selective effect on prey, and the prey develop
antipredator adaptation Anti-predator adaptations are mechanisms developed through evolution Evolution is change in the heritable Heredity, also called inheritance or biological inheritance, is the passing on of Phenotypic trait, traits from parents to thei ...
s such as
warning coloration Aposematism is the advertising Advertising is a marketing Marketing is the process of intentionally stimulating demand for and purchases of goods and services; potentially including selection of a target audience; selection of cert ...
,
alarm call "Alarm Call" is a song recorded by Icelandic singer Björk Björk Guðmundsdóttir ( ; ; born 21 November 1965) is an Icelandic singer, songwriter, record producer, actress, and DJ. Over her four-decade career, she has developed an eclect ...
s and other signals,
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard The leopard (''Pan ...

camouflage
,
mimicry In evolutionary biology Evolutionary biology is the subfield of biology that studies the evolution, evolutionary processes (natural selection, common descent, speciation) that produced the Biodiversity, diversity of life on Earth. In the 1 ...

mimicry
of well-defended species, and defensive spines and chemicals. Sometimes predator and prey find themselves in an
evolutionary arms race In evolutionary biology Evolutionary biology is the subfield of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biolo ...
, a cycle of adaptations and counter-adaptations. Predation has been a major driver of
evolution Evolution is change in the heritable Heredity, also called inheritance or biological inheritance, is the passing on of Phenotypic trait, traits from parents to their offspring; either through asexual reproduction or sexual reproduction, ...

evolution
since at least the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period A geological period is one of the several subdivisions of geologic time enabling cross-referencing of rocks and geologic events from place to place. These perio ...
period.


Definition

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; and some relationships that result in the prey's death are not generally called predation. A
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable stra ...
, such as an
ichneumon wasp
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 Parasitism is a close relationship between species In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, ...

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. There are other difficult and borderline cases.
Micropredator Parasitism is a Symbiosis, symbiotic biological interactions, relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is adaptation (biology), ad ...
s are small animals that, like predators, feed entirely on other organisms; they include
flea Flea, the common name for the order Order or ORDER or Orders may refer to: * Orderliness Orderliness is associated with other qualities such as cleanliness Cleanliness is both the abstract state of being clean and free from germs, dirt, tra ...
s and
mosquito Mosquitoes are members of a group of almost 3,600 species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defin ...

mosquito
es that consume blood from living animals, and
aphid Aphids are small sap-sucking insects and members of the Taxonomic rank, superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white Erios ...

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. Animals that on
phytoplankton Phytoplankton () are the autotrophic An autotroph or primary producer is an organism that produces complex organic compound , CH4; is among the simplest organic compounds. In chemistry, organic compounds are generally any chemical compoun ...

phytoplankton
or mats of microbes are predators, as they consume and kill their food organisms; but herbivores that browse leaves are not, as their food plants usually survive the assault. When animals eat seeds (''
seed predation Image:Aardbei muizenschade.jpg, A strawberry aggregate fruit, aggregate accessory fruit damaged by a mouse eating the seeds (achenes). Seed predation, often referred to as granivory, is a type of plant-animal interaction in which granivores (seed ...
'' or ''granivory'') or eggs (''
egg predation Egg predation is a feeding strategy in many groups of animals (ovivores) in which they consume eggs. Since an egg represents a complete organism at one stage of its Biological life cycle, life cycle, eating an egg is a form of predation, the killin ...
''), they are consuming entire living organisms, which by definition makes them predators.
Scavenger Scavengers are animals that consume dead organisms that have died from causes other than predation Predation is a biological interaction In ecology Ecology (from el, οἶκος, "house" and el, -λογία, label=none, "study ...
s, organisms that only eat organisms found already dead, are not predators, but many predators such as the
jackal Jackals are medium-sized omnivorous mammals of the Canina (subtribe), subtribe Canina, which also includes wolves and the domestic dog, among other species. While the word "jackal" has historically been used for many small canines, in modern ...

jackal
and the
hyena Hyenas, or hyaenas (from Ancient Greek , ), are feliformia, feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in their own genus), it is the fifth-smallest biological family in the Carnivora and one of the ...

hyena
scavenge when the opportunity arises. Among invertebrates, social wasps (yellowjackets) are both hunters and scavengers of other insects.


Taxonomic range

While examples of predators among mammals and birds are well known, predators can be found in a broad range of taxa including arthropods. They are common among insects, including mantids,
dragonflies A dragonfly is a flying insect belonging to the order Odonata, infraorder Anisoptera (from Ancient Greek, Greek ἄνισος ''anisos'', "unequal" and πτερόν ''pteron'', "wing", because the hindwing is broader than the forewing). Adult d ...

dragonflies
,
lacewings The insect order (biology), order Neuroptera, or net-winged insects, includes the lacewings, Mantispidae, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera can be grouped together with the Megalopte ...

lacewings
and
scorpionflies Mecoptera (from the Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 ...
. In some species such as the
alderfly Alderflies are megalopteran insects of the Family (biology), family Sialidae. They are closely related to the dobsonflies and fishflies as well as to the prehistoric Euchauliodidae. All living alderflies – about 66 species all together – are ...
, only the larvae are predatory (the adults do not eat). Spiders are predatory, as well as other terrestrial invertebrates such as
scorpion Scorpions are predatory Predation is a biological interaction In ecology Ecology (from el, οἶκος, "house" and el, -λογία, label=none, "study of") is the study of the relationships between living organisms, including hum ...

scorpion
s;
centipede Centipedes (from the New Latin New Latin (also called Neo-Latin or modern Latin) is the revival of Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally s ...

centipede
s; some
mite Mites are small arachnid Arachnida () is a class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Cl ...
s,
snail A snail is, in loose terms, a shelled gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic Taxonomy (general) is the practice and science of classification of things or concepts, including the pr ...

snail
s and
slug Slug, or land slug, is a common name Common may refer to: Places * Common, a townland in County Tyrone, Northern Ireland * Boston Common Boston Common (also known as the Common) is a central public park in downtown Boston, Massachusetts. ...

slug
s;
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...

nematode
s; and planarian worms. In marine environments, most
cnidaria Pacific sea nettles, ''Chrysaora fuscescens'' Cnidaria () is a phylum In biology, a phylum (; plural The plural (sometimes list of glossing abbreviations, abbreviated ), in many languages, is one of the values of the grammatical number, ...

cnidaria
ns (e.g.,
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum In zoological nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted Con ...

jellyfish
, hydroids),
ctenophora Ctenophora (; ctenophore ; ) comprise a phylum of marine life, marine invertebrates, commonly known as comb jellies, that marine habitats, inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly r ...

ctenophora
(comb jellies),
echinoderm An echinoderm () is any member of the phylum In biology, a phylum (; plural The plural (sometimes list of glossing abbreviations, abbreviated ), in many languages, is one of the values of the grammatical number, grammatical category of ...
s (e.g.,
sea stars Starfish or sea stars are star-shaped echinoderm An echinoderm is any member of the phylum In biology, a phylum (; plural The plural (sometimes list of glossing abbreviations, abbreviated ), in many languages, is one of the values ...
,
sea urchin Sea urchins () are spine (zoology), spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone — from the intertidal seashore down to . The spherical, ha ...

sea urchin
s, sand dollars, and
sea cucumber Sea cucumbers are echinoderms from the class (biology), class Holothuroidea. They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the sea floor worldwide. The number o ...

sea cucumber
s) and
flatworm The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a count ...

flatworm
s are predatory. Among
crustacean Crustaceans (Crustacea ) form a large, diverse arthropod taxon which includes such animals as crabs, lobsters, crayfish, Caridea, shrimp, krill, Dendrobranchiata, prawns, woodlice, barnacles, copepods, amphipoda, amphipods and mantis shrimp. The ...
s,
lobster Lobsters are a family In , family (from la, familia) is a of people related either by (by recognized birth) or (by marriage or other relationship). The purpose of families is to maintain the well-being of its members and of society ...

lobster
s,
crab Crabs are decapod crustacean Crustaceans (Crustacea ) form a large, diverse arthropod taxon which includes such animals as crabs, lobsters, crayfish, Caridea, shrimp, krill, Dendrobranchiata, prawns, woodlice, barnacles, copepods, amphipoda ...

crab
s,
shrimp Shrimp are Decapoda, decapod crustaceans with elongated bodies and a primarily swimming mode of locomotion – most commonly Caridea and Dendrobranchiata. More narrow definitions may be restricted to Caridea, to smaller species of either group ...

shrimp
s and
barnacle A barnacle is a type of arthropod An arthropod (, (gen. ποδός)) is an invertebrate animal having an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Euarthropoda,Reference ...

barnacle
s are predators, and in turn crustaceans are preyed on by nearly all
cephalopod A cephalopod is any member of the mollusca Mollusca is the second-largest phylum In biology, a phylum (; plural The plural (sometimes list of glossing abbreviations, abbreviated ), in many languages, is one of the values of the gram ...
s (including
octopus An octopus (pl. octopuses/octopi, see below for variants) is a soft-bodied, eight- limbed mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda. The members are known as molluscs or mollusks (). A ...

octopus
es,
squid Squid are cephalopod A cephalopod is any member of the mollusca Mollusca is the second-largest phylum In biology, a phylum (; plural The plural (sometimes list of glossing abbreviations, abbreviated ), in many languages, is ...

squid
and
cuttlefish Cuttlefish or cuttles are marine Marine is an adjective meaning of or pertaining to the sea or ocean The ocean (also the sea or the world ocean) is the body of salt water which covers approximately 71% of the surface of the Earth.
cuttlefish
). Seed predation is restricted to mammals, birds, and insects but is found in almost all terrestrial ecosystems. Egg predation includes both specialist egg predators such as some
colubrid Colubridae (, commonly known as colubrids , from la, coluber, 'snake') is a family In human society, family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other ...
snake Snakes are elongated, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Many species of snakes ...

snake
s and generalists such as foxes and badgers that opportunistically take eggs when they find them. Some plants, like the
pitcher plant 250px, Scanning electron micrograph of a pitcher's inner surface Pitcher plants are several different carnivorous plants which have modified leaves known as pitfall traps—a prey-trapping mechanism featuring a deep cavity filled with digesti ...

pitcher plant
, the and the
sundew ''Drosera'', which is commonly known as the sundews, is one of the largest genera Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology, taxonomy () is the scientific s ...

sundew
, are . 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. Some carnivorous fungi catch
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...

nematode
s using either active traps in the form of constricting rings, or passive traps with adhesive structures. Many species of
protozoa Protozoa (singular protozoon or protozoan, plural protozoa or protozoans) is an informal term for a group of single-celled eukaryote Eukaryotes () are organism In biology, an organism () is any organic, life, living system that ...

protozoa
(
eukaryote Eukaryotes () are organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interact ...

eukaryote
s) and
bacteria Bacteria (; common noun bacteria, singular bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typ ...

bacteria
(
prokaryote A prokaryote () is a single-celled organism A unicellular organism, also known as a single-celled organism, is an organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual conti ...
s) prey on other microorganisms; the feeding mode is evidently ancient, and evolved many times in both groups. Among freshwater and marine
zooplankton Zooplankton (; ) are heterotroph A heterotroph (; from Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often ro ...

zooplankton
, whether single-celled or multi-cellular, predatory grazing on
phytoplankton Phytoplankton () are the autotrophic An autotroph or primary producer is an organism that produces complex organic compound , CH4; is among the simplest organic compounds. In chemistry, organic compounds are generally any chemical compoun ...

phytoplankton
and smaller zooplankton is common, and found in many species of nanoflagellates,
dinoflagellate The dinoflagellates (Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 m ...
s,
ciliate The ciliates are a group of protozoan Protozoa (singular protozoon or protozoan, plural protozoa or protozoans) is an informal term for a group of single-celled eukaryote Eukaryotes () are organisms whose Cell (biology), cells have a c ...

ciliate
s,
rotifer The rotifers (, from Latin ''wikt:rota#Latin, rota'' "wheel" and ''wikt:-fer#Latin, -fer'' "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic Coelom#Pseudocoelomates, ...

rotifer
s, a diverse range of
meroplankton Meroplankton are a wide variety of aquatic organisms which have both planktonic and benthic stages in their life cycles. Much of the meroplankton consists of larval stages of larger organism. Meroplankton can be contrasted with holoplankton, w ...
animal larvae, and two groups of crustaceans, namely
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor), ...

copepod
s and
cladocera The Cladocera are an order of small crustaceans commonly called water fleas. Over 650 species have been recognised so far, with many more undescribed. They first appeared before the Permian The Permian ( ) is a geologic period and stratigra ...
ns. summarizes findings from many authors.


Foraging

To feed, a predator must search for, pursue and kill its prey. These actions form a
foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology Behavioral ecology, also spe ...
cycle. 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). 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).


Search

Predators have a choice of search modes ranging from ''sit-and-wait'' to ''active'' or ''widely foraging''. The sit-and-wait method is most suitable if the prey are dense and mobile, and the predator has low energy requirements. Wide foraging expends more energy, and is used when prey is sedentary or sparsely distributed. There is a continuum of search modes with intervals between periods of movement ranging from seconds to months. Sharks, sunfish,
Insectivorous A robber fly eating a _.html" ;"title="hoverfly ">hoverfly An insectivore is a Carnivore">carnivorous A carnivore , meaning "meat Meat is animal flesh that is eaten as food. Humans have hunted and killed animals for meat since pre ...
birds and
shrew Shrews (family In human society A society is a Social group, group of individuals involved in persistent Social relation, social interaction, or a large social group sharing the same spatial or social territory, typically subject to ...
s are almost always moving while web-building spiders, aquatic invertebrates, praying mantises and
kestrel The name kestrel (from french: crécerelle, derivative from , i.e. ratchet) is given to several members of the falcon genus, ''Falco''. Kestrels are most easily distinguished by their typical hunting behaviour which is to hover at a height of ar ...

kestrel
s rarely move. In between,
plover Plovers ( or ) are a widely distributed group of wader, wading birds belonging to the subfamily Charadriinae. Description There are about 66 species in the subfamily, most of them called "plover" or "Dotterel (disambiguation), dotterel". Th ...
s and other
shorebirds Waders or shorebirds are bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves , characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high M ...
, freshwater fish including crappies, and the larvae of Coccinellidae, coccinellid beetles (ladybirds), alternate between actively searching and scanning the environment. Prey distributions are often clumped, and predators respond by looking for ''patches'' where prey is dense and then searching within patches. 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. For example, the black-browed albatross regularly makes foraging flights to a range of around , up to a maximum foraging range of for breeding birds gathering food for their young. With static prey, some predators can learn suitable patch locations and return to them at intervals to feed. The optimal foraging theory, optimal foraging strategy for search has been modelled using the marginal value theorem. 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.


Assessment

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. 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. A predator may also assess a patch and decide whether to spend time searching for prey in it. This may involve some knowledge of the preferences of the prey; for example, ladybirds can choose a patch of vegetation suitable for their
aphid Aphids are small sap-sucking insects and members of the Taxonomic rank, superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white Erios ...

aphid
prey.


Capture

To capture prey, predators have a spectrum of pursuit modes that range from overt chase (''
pursuit predation Pursuit predation is a form of predation Predation is a biological interaction In ecology Ecology (from el, οἶκος, "house" and el, -λογία, label=none, "study of") is the study of the relationships between living organisms ...
'') to a sudden strike on nearby prey (''ambush predator, ambush predation''). 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.


Ambush

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. Vertebrate ambush predators include frogs, fish such as the angel shark, the northern pike and the eastern frogfish. Among the many invertebrate ambush predators are trapdoor spiders and Thomisus spectabilis, Australian Crab spiders on land and mantis shrimps in the sea. 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. The capturing movement has to be rapid to trap the prey, given that the attack is not modifiable once launched.


Ballistic interception

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. 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. Ballistic predators include insects such as dragonflies, and vertebrates such as archerfish (attacking with a jet of water), chameleons (attacking with their tongues), and some Colubridae, colubrid snakes.


Pursuit

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. 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. Many pursuit predators use camouflage to approach the prey as close as possible unobserved (''stalking'') before starting the pursuit. 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; predatory birds (raptors) such as falcons; and insects such as dragonfly, dragonflies. 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-gatherers and by Canidae, canids such as African wild dogs 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. A specialised form of pursuit predation is the lunge feeding of baleen whales. 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 filtering it through their feathery baleen plates. Pursuit predators may be Social predator, social, like the lion and wolf that hunt in groups, or solitary.


Handling

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.


Solitary versus social predation

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 Hyenas, or hyaenas (from Ancient Greek , ), are feliformia, feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in their own genus), it is the fifth-smallest biological family in the Carnivora and one of the ...

hyena
s, and Wolf, wolves collaborate to catch and kill herbivores as large as buffalo, and lions even hunt elephants. 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 dolphins form a circle around a school of fish and move inwards, concentrating the fish by a factor of 200. By hunting socially Common chimpanzee, chimpanzees can catch colobus monkeys that would readily escape an individual hunter, while cooperating Harris hawks can trap rabbits. 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
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es. These predators are able to access small crevices and flush out the prey. Killer whales have been known to help whalers hunt baleen whales. ISBN R-105732-9. 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. Ambush predators are often solitary to reduce the risk of becoming prey themselves. 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, including the cougar and cheetah. However, the solitary cougar does allow other cougars to share in a kill, and the coyote can be either solitary or social. Other solitary predators include the northern pike, wolf spiders and all the thousands of species of solitary wasps among arthropods, and many microorganisms and
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.


Specialization


Physical adaptations

Under the pressure of natural selection, predators have evolved a variety of physical adaptations for detecting, catching, killing, and digesting prey. These include speed, agility, stealth, sharp senses, claws, teeth, filters, and suitable digestive systems. For Prey detection, detecting prey, predators have well-developed
vision Vision or The Vision may refer to: Perception Optical perception * Visual perception Visual perception is the ability to interpret the surrounding environment (biophysical), environment through photopic vision (daytime vision), color visio ...

vision
, smell, or
hearing Schematic diagram of the human ear Hearing, or auditory perception, is the ability to perceive Sound, sounds through an organ, such as an ear, by detecting Vibration, vibrations as periodic changes in the pressure of a surrounding medium. Th ...

hearing
. Predators as diverse as owls and jumping spiders 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 of snow or earth. Many predators have acute hearing, and some such as animal echolocation, echolocating bats hunt exclusively by active or passive use of sound. Predators including big cats, 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 Snakes are elongated, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Many species of snakes ...

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. Fish and other predators have developed the ability to crush or open the armoured shells of molluscs. Many predators are powerfully built and can catch and kill animals larger than themselves; this applies as much to small predators such as ants and
shrew Shrews (family In human society A society is a Social group, group of individuals involved in persistent Social relation, social interaction, or a large social group sharing the same spatial or social territory, typically subject to ...
s as to big and visibly muscular carnivores like the cougar and lion. 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 eyes, 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


Diet and behaviour

Predators are often highly specialized in their diet and hunting behaviour; for example, the Eurasian lynx only hunts small ungulates. Others such as leopards are more opportunistic generalists, preying on at least 100 species. 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; this appears to be correct for predators of immobile prey, but is doubtful with mobile prey. In size-selective predation, predators select prey of a certain size. 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.


Camouflage and mimicry

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 patterns suiting their habitats. In
aggressive mimicry Aggressive mimicry is a form of mimicry In evolutionary biology Evolutionary biology is the subfield of biology that studies the evolution, evolutionary processes (natural selection, common descent, speciation) that produced the Biod ...
, 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. Flower mantises are ambush predators; camouflaged as flowers, such as orchids, they attract prey and seize it when it is close enough. Frogfishes are extremely well camouflaged, and actively lure their prey to approach using an 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.


Venom

Many smaller predators such as the box jellyfish use venom to subdue their prey, and venom can also aid in digestion (as is the case for rattlesnakes and some spiders). 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 nucleotides) that inactives it. These changes are explained by the fact that its prey does not need to be subdued.


Electric fields

Several groups of predatory fish have the ability to detect, track, and sometimes, as in the electric ray, to incapacitate their prey by generating electric fields using Electric organ (biology), electric organs. The electric organ is derived from modified nerve or muscle tissue.


Physiology

Physiological adaptations to predation include the ability of predatory bacteria to digest the complex peptidoglycan polymer from the cell walls of the bacteria that they prey upon. 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 proteins in their diet.


Antipredator adaptations

To counter predation, prey have evolved defences for use at each stage of an attack. They can try to avoid detection, such as by using
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard The leopard (''Pan ...

camouflage
and
mimicry In evolutionary biology Evolutionary biology is the subfield of biology that studies the evolution, evolutionary processes (natural selection, common descent, speciation) that produced the Biodiversity, diversity of life on Earth. In the 1 ...

mimicry
. They can detect predators and warn others of their presence. If detected, they can try to avoid being the target of an attack, for example, by Aposematism, signalling that they are toxic or unpalatable, by forming groups, or that a chase would be unprofitable. If they become a target, they can try to fend off the attack with defences such as armour, Spine (zoology), quills, unpalatability, or mobbing; and they can escape an attack in progress by deimatic behaviour, startling the predator, Thanatosis, playing dead, shedding body parts such as tails, or simply fleeing.


Coevolution

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. Many pursuit predators that run on land, such as wolves, have evolved long limbs in response to the increased speed of their prey. Their adaptations have been characterized as an
evolutionary arms race In evolutionary biology Evolutionary biology is the subfield of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biolo ...
, an example of the coevolution of two species. 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. 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. 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, 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. 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. Also, if a predator loses enough dinners, it too will lose its life. 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. 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. 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. Their hearing evolved in response to bat predation, but the only clear example of reciprocal adaptation in bats is stealth echolocation. 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.


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.


Trophic level

One way of classifying predators is by trophic level. Carnivores that feed on herbivores are secondary consumers; their predators are tertiary consumers, and so forth. At the top of this food chain are apex predators such as lions. Many predators however eat from multiple levels of the food chain; a carnivore may eat both secondary and tertiary consumers. This means that many predators must contend with intraguild predation, where other predators kill and eat them. For example, coyotes compete with and sometimes kill gray foxes and bobcats.


Biodiversity maintained by apex predation

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. 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. 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 beavers' food sources. The removal of wolves had a direct effect on the beaver population, as their habitat became territory for grazing. Increased browsing on willows and conifers 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.


Population dynamics

In the absence of predators, the population of a species can grow exponentially until it approaches the carrying capacity of the environment. Predators limit the growth of prey both by consuming them and by changing their behavior. 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 forests 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 hunters for the Hudson Bay Company over more than a century. A simple model of a system with one species each of predator and prey, the Lotka–Volterra equations, predicts population cycles. 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. 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. 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. 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. Many factors can stabilize predator and prey populations. 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. As a result, population cycles tend to be found in northern temperate and subarctic ecosystems because the food webs are simpler. The snowshoe hare-lynx system is subarctic, but even this involves other predators, including coyotes, goshawks and great horned owls, and the cycle is reinforced by variations in the food available to the hares. 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 distributions, or to animal migration, migrate; to have differences between individuals, such as sexes and an age structure, so that only some individuals reproduce; to live in a varying environment, such as with changing seasons; 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. 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.


Evolutionary history

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. 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. It likely triggered major evolutionary transitions including the arrival of Cell (biology), cells,
eukaryote Eukaryotes () are organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interact ...

eukaryote
s, sexual reproduction, multicellularity, increased size, mobility (including insect flight) and armoured shells and exoskeletons. 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). Predation visibly became important shortly before the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period A geological period is one of the several subdivisions of geologic time enabling cross-referencing of rocks and geologic events from place to place. These perio ...
period—around —as evidenced by the almost simultaneous development of calcification in animals and algae, and predation-avoiding burrowing. However, predators had been grazing on micro-organisms since at least , with evidence of selective (rather than random) predation from a similar time. 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. Among the Cambrian predators were invertebrates like the anomalocaridids with appendages suitable for grabbing prey, large compound eyes and jaws made of a hard material like that in the exoskeleton of an insect. 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 ''Dunkleosteus'', is considered the world's first
vertebrate Vertebrates () comprise all species of animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic ma ...
"superpredator", preying upon other predators. Insects developed the ability to fly in the Early Carboniferous or Late Devonian, enabling them among other things to escape from predators. Among the largest predators that have ever lived were the theropod dinosaurs such as ''Tyrannosaurus'' from the Cretaceous period. They preyed upon herbivorous dinosaurs such as hadrosaurs, ceratopsians and ankylosaurs. 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 The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period A geological period is one of the several subdivisions of geologic time enabling cross-referencing of rocks and geologic events from place to place. These perio ...
predators. File:Laggania cambria 02.jpg, Mouth of the anomalocaridid ''Laggania cambria'', a
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period A geological period is one of the several subdivisions of geologic time enabling cross-referencing of rocks and geologic events from place to place. These perio ...
invertebrate, probably an apex predator File:Dunkleosteus terrelli - MUSE.jpg, ''Dunkleosteus'', a Devonian placoderm, perhaps the world's first vertebrate superpredator, reconstruction File:Meganeura monyi au Museum de Toulouse.jpg, ''Meganeura monyi'', a predatory Carboniferous insect related to
dragonflies A dragonfly is a flying insect belonging to the order Odonata, infraorder Anisoptera (from Ancient Greek, Greek ἄνισος ''anisos'', "unequal" and πτερόν ''pteron'', "wing", because the hindwing is broader than the forewing). Adult d ...

dragonflies
, could fly to escape terrestrial predators. Its large size, with a wingspan of , may reflect the lack of vertebrate aerial predators at that time. File:Tyrannosaurus model at NHM.jpg, ''Tyrannosaurus'', a large theropod dinosaur of the Cretaceous, reconstruction


In human society


Practical uses

Humans, as omnivores, are to some extent predatory, using weapons and tools to fishing, fish, hunting, hunt and animal trapping, trap animals. They also use other predatory species such as Hunting dog, dogs, cormorants, and falcons to catch prey for food or for sport. Two mid-sized predators, dogs and cats, are the animals most often kept as pets in western societies. 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. 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 pesticides.


Symbolic uses

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. A terrifying predator, a gigantic Man-eater, man-eating great white shark, is central, too, to Steven Spielberg's 1974 thriller ''Jaws (film), Jaws''. Among poetry on the theme of predation, a predator's consciousness might be explored, such as in Ted Hughes's ''Pike''. 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. In mythology and folk fable, predators such as the fox and wolf have mixed reputations. translated from Wallner, A. (1998) ''Die Bedeutung der Raubtiere in der Mythologie: Ergebnisse einer Literaturstudie''. - Inf.bl. Forsch.bereiches Landsch.ökol. 39: 4-5. 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. 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. In the Middle Ages, belief spread in werewolf, werewolves, men transformed into wolves. 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. More recently, in Rudyard Kipling's 1894 ''The Jungle Book'', Mowgli is raised by the wolf pack. 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.


See also

* Ecology of fear (concept), Ecology of fear * Predation problem * Predator–prey reversal * Wa-Tor


Notes


References


Sources

* * * * * * *


External links

* * {{good article Ecology Predation, Biological pest control