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Solitary predator: a polar bear feeds on a bearded seal it has killed
Social predators: meat ants cooperate to feed on a cicada far larger than themselves

Predation is a biological interaction where a predator (an organism, often an animal) kills and eats its prey (another organism). Predators are adapted and often highly specialized for hunting, with acute vision, hearing, and sense of smell. Many predatory animals, both vertebrates such as lions and sharks, and invertebrates such as mantises, have sharp claws or jaws to grip, kill, and cut up their prey. When an animal's prey is passive, as with seed and egg predators, however, such adaptations are often reduced or absent.

In ecology, predators are heterotrophic, getting all their energy from other organisms. This places them at high trophic levels in food webs. Many predators are carnivores, an exception being seed predators. Predation is one of a family of common feeding behaviours that includes parasitism and micropredation which usually do not kill the host, and parasitoidism which always does, eventually.

Predator and prey adapt to each other in an evolutionary arms race, coevolving under natural selection to develop antipredator adaptations such as camouflage and defensive spines and chemicals in the prey, and adaptations such as stealth and aggressive mimicry that improve hunting efficiency in the predator. Predation has a powerful selective effect, especially on prey, and it has been a major driver of evolution since at least the Cambrian period.

Contents

DefinitionEdit

 
Similar strategies: spider wasps paralyse and eventually kill their hosts, but they are considered parasitoids 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; and some relationships that result in the prey's death are not generally 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 two ways: it kills its prey immediately; and 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.[1][2]

The boundary with scavenging, eating the bodies of animals found already dead, is likewise blurred, since many predators such as the jackal and the hyena scavenge when the opportunity arises.[3][4] Among invertebrates, social wasps (yellowjackets) are both hunters and scavengers of other insects.[5]

 
Diagram of the boundaries and near neighbours of predation as a feeding strategy

There are other difficult and borderline cases. For example, micropredators are small animals that, like predators, feed entirely on other organisms; they include fleas and mosquitoes that consume blood from living animals, and aphids that consume sap from living plants. However, since micropredators typically do not kill their hosts, they are now often thought of as parasites.[6][7] As another example, animals that eat plants are generally thought of as non-predatory herbivores, contrasted with predatory carnivores, but when those animals eat seeds (seed predation or granivory) or eggs (egg predation), they are consuming entire living organisms, which by definition makes them predators.[8][9][10][8] Many predators are also scavengers, but pure scavengers that eat only dead organisms are not predators.[11] Animals that graze on 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.[11] In the words of the paleontologist Stefan Bengtson, predation "involves much more than fanged beasts that pounce with a roar upon the hapless leaf-muncher."[11]

Taxonomic rangeEdit

Carnivorous plant: sundew engulfing an insect
Seed predation: mouse eating seeds

While examples of predators among mammals and birds are well known,[12] predators can be found in a broad range of taxa. They are common among insects, including mantids, dragonflies, lacewings and 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 scorpions; centipedes; some mites, snails and slugs; nematodes; and planarian worms.[13] In marine environments, most cnidarians (e.g., jellyfish, hydroids), ctenophora (comb jellies), echinoderms (e.g., sea stars, sea urchins, sand dollars, and sea cucumbers) and flatworms are predatory.[14] Among crustaceans, lobsters, crabs, shrimps and barnacles are predators,[15] and in turn crustaceans are preyed on by nearly all cephalopods (including octopuses, squid and cuttlefish).[16]

 
Paramecium, a predatory ciliate, feeding on bacteria

Seed predation is restricted to mammals, birds, and insects and is found in almost all terrestrial ecosystems.[10][8] Egg predation includes both specialist egg predators such as some colubrid snakes and generalists such as foxes and badgers that opportunistically take eggs when they find them.[17][18][19]

Some plants, like the pitcher plant, the Venus fly trap and the sundew, are carnivorous and consume insects.[12] Some carnivorous fungi catch nematodes using either active traps in the form of constricting rings, or passive traps with adhesive structures.[20]

Many species of protozoa (eukaryotes) and bacteria (prokaryotes) prey on other microorganisms; the feeding mode is evidently ancient, and evolved many times in both groups.[21][12][22] 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 nanoflagellates, dinoflagellates, ciliates, rotifers, a diverse range of meroplankton animal larvae, and two groups of crustaceans, namely copepods and cladocerans.[23]

StrategiesEdit

 
A basic foraging sequence for a predator, with some variations indicated[24]

Overview: foraging modesEdit

Predators have a spectrum of foraging modes that range from hunting actively for prey (pursuit predation) to sitting and waiting for prey to approach within striking distance (ambush predation).[25][12] Closely related to pursuit is stalking where a predator stealthily searches for prey and then pursues it over a short distance.[26][27] 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.[28] In all foraging modes, there is a sequence of stages, which can be described broadly (with many variations) as search, assessment, pursuit or its equivalent, and handling.[24]

Pursuit predators may be social, like the lion and wolf that hunt in groups, or solitary, like the cheetah.[2]

Pursuit predationEdit

Humpback whales are lunge feeders, filtering thousands of krill from seawater and swallowing them alive.
Dragonflies, like this common clubtail with captured prey, are invertebrate pursuit predators.

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.[28] 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.[28] Many pursuit predators use camouflage to approach the prey as close as possible unobserved (stalking) before starting the pursuit.[28] Pursuit predators include terrestrial mammals such as lions, cheetahs, and wolves; marine predators such as dolphins and many predatory fishes, such as tuna;[29][30] predatory birds (raptors) such as falcons; and insects such as dragonflies.[31]

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.[32][33]

Ambush predationEdit

A trapdoor spider waiting in its burrow to ambush its prey

Ambush or sit-and-wait predators are carnivorous animals or other organisms, such as some nematophagous fungi and carnivorous plants, 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.[34][28] Vertebrate ambush predators include frogs, fish such as the angel shark, the northern pike and the eastern frogfish.[28][35][36][37] Among the many invertebrate ambush predators are trapdoor spiders on land and mantis shrimps in the sea.[34][38][39] 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.[28] The capturing movement has to be rapid to trap the prey, given that the attack is not modifiable once launched.[28]

Ballistic interceptionEdit

 
Chameleon attacks prey by shooting out its 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.[28] 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.[28] Ballistic predators include insects such as dragonflies, vertebrates such as archerfish (attacking with a jet of water), chameleons (attacking with their tongues), and some colubrid snakes.[28]

Solitary versus social predationEdit

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, hyenas, and wolves collaborate to catch and kill herbivores as large as buffalo, and lions even hunt elephants.[40][41][42] 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.[43] By hunting socially chimpanzees can catch colobus monkeys that would readily escape an individual hunter, while cooperating Harris hawks can trap rabbits.[40][44]

 
Wolves, social predators, cooperate to hunt and kill bison.

Cooperative behavior can occur between predators of different species. In coral reefs, when fish such as the grouper and coral trout spot prey that is inaccessible to them, they signal to giant moray eels, Napoleon wrasses or octopuses. These predators are able to access small crevices and flush out the prey.[45][46] Killer whales have been known to help whalers hunt baleen whales.[47]

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.[48][49] Ambush predators are often solitary to reduce the risk of becoming prey themselves.[50] Of 245 terrestrial carnivores, 177 are solitary; and 35 of the 37 wild cats are solitary,[51] including the cougar and cheetah.[48][2] However, the solitary cougar does allow other cougars to share in a kill,[52] and the coyote can be either solitary or social.[53] Other solitary predators include the northern pike,[54] wolf spiders and solitary wasps,[55][56] and many microorganisms and zooplankton.[21][57]

SpecializationEdit

Physical adaptationsEdit

Under the pressure of natural selection, predators have evolved a variety of physical adaptations for detecting, catching, killing, and digesting prey. For detecting prey, predators have well-developed vision, smell, or hearing.[12] 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 2 feet (60 cm) of snow or earth. Many predators have acute hearing, and some such as echolocating bats hunt exclusively by active or passive use of sound.[58]

Predators including big cats, 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 snakes 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.[58]

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 shrews as to big and visibly muscular carnivores like the cougar and lion.[58][2][59]

Diet and behaviourEdit

 
Platydemus manokwari, a specialist flatworm predator of land snails

Predators are often highly specialized in their diet and hunting behaviour; for example, the Eurasian lynx only hunts small ungulates.[60] Others such as leopards are more opportunistic generalists, preying on at least 100 species.[61][62] 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;[63] this appears to be correct for predators of immobile prey, but is doubtful with mobile prey.[64]

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 for large prey. For example, adult elephants are relatively safe from predation by lions, but juveniles are vulnerable.[65][66][67] Many smaller predators such as the box jellyfish use venom to subdue their prey,[68] and venom can also aid in digestion (as is the case for rattlesnakes and some spiders).[69][70]

PhysiologyEdit

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.[22] 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.[71]

Coevolution with preyEdit

The relationships between predators and their prey are at the heart of ecosystem dynamics. These have traditionally been explored as feeding and energy flows;[58] more recently, they have been viewed as interactions of the physiology, morphology, and behaviour of predator and prey. For example, a predator may evolve to become more aggressive, while the prey may become more alert; a predator could evolve a wider gape of its jaws to accommodate larger prey, while the prey may become larger and hence more difficult to kill. Such effects often interact in multiple ways.[26]

Antipredator adaptationsEdit

Springbok stotting to signal its ability to escape
Dead leaf mantis's camouflage makes it less visible to both predators and prey.

A hunting predator is attempting to obtain its next meal; its prey is attempting to save its own life. This sets up an evolutionary arms race, causing many antipredator adaptations to evolve in prey populations due to the selective pressures of predation over long periods of time.[12][72][73]

Many prey animals are aposematically coloured or patterned as a warning to predators that they are distasteful or able to defend themselves.[58][74][75] Such distastefulness or toxicity is brought about by chemical defences, found in a wide range of prey, especially insects, but the skunk is a dramatic mammalian example.[76] Chemical defences include toxins, such as bitter compounds in leaves absorbed by leaf-eating insects and used to dissuade potential predators.[77] Mechanical defences include sharp spines, hard shells and tough leathery skin or exoskeletons, all making prey harder to kill.[78]

Some species mob predators cooperatively, reducing the likelihood of attack.[79] Others such as Thomson's gazelle stot to signal to predators such as cheetahs that they will have an unprofitable chase.[80]

Camouflage makes use of coloration, shape, and pattern to misdirect the visual sensory mechanisms of predators, enabling prey to remain unrecognized.[58][81] Among the many mechanisms of camouflage are countershading[82] and disruptive coloration.[83] The resemblance can be to the biotic or non-living environment, such as a mantis resembling dead leaves, or to other organisms. In mimicry, an organism has a similar appearance to another species, as in the drone fly, which resembles a bee, yet has no sting. Many butterflies and moths have eyespots, wing markings that resemble eyes.[84] When a predator disturbs the insect, it reveals its hind wings, startling the predator and giving it time to escape.[85][86]

Antiprey adaptationsEdit

Striated frogfish uses camouflage and aggressive mimicry in the form of a fishing rod-like lure on its head to attract prey.

As prey evolves to become harder to catch, on their side of the evolutionary arms race predators adapt to use speed, stealth, camouflage and aggressive mimicry to improve their hunting efficiency, an example of coevolution. For example, many pursuit predators that run on land, such as wolves, have evolved long limbs in response to the increased speed of their prey.[87]

Members of the cat family such as the snow leopard (treeless highlands), tiger (grassy plains, reed swamps), ocelot (forest), fishing cat (waterside thickets), and lion (open plains) have coloration and patterns suiting their habitats.[88] Female Photuris fireflies, for example, copy the light signals of other species, thereby attracting male fireflies, which they capture and eat.[89] Flower mantises are ambush predators; camouflaged as flowers, such as orchids, they attract prey and seize it when it is close enough.[90] 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.[91] Predators have evolved whatever capabilities have helped them to catch and eat their prey, including speed, agility, stealth, sharp senses, claws, teeth, filters, and suitable digestive systems.[92] For example, fish and other predators have developed the ability to crush or open the armoured shells of molluscs.[93]

 
Predation as competition: a bobcat and two coyotes facing off over a kill. Coyotes sometimes kill smaller predators including bobcats.

As competitionEdit

An alternative view considers predation as a form of competition with the prey: the predator's and prey's genes are competing for the prey's body.[94]

Competition is in addition directly evident in intraguild predation, where predators kill and eat predators of competing species at the same trophic level. For example, coyotes compete with and sometimes kill gray foxes and bobcats.[95]

Role in ecosystemsEdit

Trophic levelEdit

 
Secondary consumer: a mantis (Tenodera aridifolia) eating a bee

Predators are often another organism's prey, and likewise prey are often predators. Though blue jays prey on insects, they may in turn be prey for cats and snakes, and snakes may be the prey of hawks. One way of classifying predators is by trophic level. Carnivores that feed on heterotrophs are secondary consumers; their predators are tertiary consumers, and so forth. Because only a fraction of energy is passed on to the next level, this hierarchy of predation must end somewhere, and very seldom goes higher than five or six levels.[96]

For example, a lion, an apex predator (at the top of its food chain, if parasites are not considered) that preys upon large herbivores such as wildebeest, which in turn eat grasses, is only a secondary consumer. Other apex predators include the sperm whale, Komodo dragon, tiger, and most eagles and owls.[96]

Many predators eat from multiple levels of the food chain. A carnivore may eat both secondary and tertiary consumers, and its prey may itself be difficult to classify for similar reasons.[96]

Biodiversity maintained by apex predationEdit

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.[97] 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.[98]

 
Riparian willow recovery at Blacktail Creek, Yellowstone National Park, after reintroduction of wolves, the local keystone species and apex predator.[99] 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 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.[99]

Population dynamicsEdit

 
Lotka–Volterra equations model population dynamics of predator–prey interactions, creating linked cycles in the two populations.

Predators tend to lower the survival and fecundity of their prey, but also depend on prey for their survival, so predator populations are affected by changes in prey populations and vice versa. The population dynamics of predator–prey interactions can be modeled using the Lotka–Volterra equations, dating from the early 20th century. These provide a mathematical model for the cycling of predator and prey populations.[100] Predators tend to select young, weak, and ill individuals, leaving prey populations able to regrow. When prey numbers are low, the predators find little food, produce few young, and may starve, so their population tends to fall. When predator numbers are low, few prey are killed, and they can reproduce freely, so their population grows. Once prey numbers are high, the predators catch food more readily and raise more young, so their population grows. When predator and prey numbers are high, the predators kill many prey, depleting their population, and their numbers fall. The cycles then repeat.[101][102]

Evolutionary historyEdit

Predation have evolved repeatedly in different groups of organisms. When predation appears in an ecosystem, it becomes a powerful evolutionary force—since every successful predation event means the death of a prey organism,[a] driving the coevolution of changes in prey and predators. Predation, too, likely triggered major evolutionary transitions including the arrival of cells, eukaryotes, sexual reproduction, multicellularity, increased size, mobility, and armoured shells and exoskeletons.[11]

Predation visibly became important shortly before the Cambrian period—around 550 million years ago—as evidenced by the almost simultaneous development of calcification in animals and algae,[103] and predation-avoiding burrowing. However, predators had been grazing on micro-organisms since at least 1,000 million years ago,[11][104][105][106] with evidence of selective (rather than random) predation from a similar time.[107]

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.[108] Among the Cambrian predators were invertebrates like the anomalocaridids, euarthropods with raptorial appendages, large compound eyes and hard sclerotised jaws.[109] Some of the first fish to have jaws were the armoured and mainly predatory placoderms of the Silurian to Devonian periods; the 6 m (20 ft) placoderm Dunkleosteus is considered the world's first vertebrate "superpredator", preying upon other predators.[110][111]Insects developed the ability to fly in the Early Carboniferous or Late Devonian, enabling them among other things to escape from predators.[112] 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.[113]

In human societyEdit

 
San hunter, Botswana

Humans are to some extent predatory,[114] fishing,[115] hunting and trapping animals using weapons and tools.[116] They also use other predatory species, such as dogs, cormorants,[117] and falcons to catch prey for food or for sport.[118] Neolithic hunters, including the San of southern Africa, used persistence hunting, a form of pursuit predation where the pursuer may be slower than the prey over short distances, to exhaust the prey such as a kudu antelope in the midday heat over a period of up to five hours.[119][120]

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 not do harm to non-pest species, are an environmentally friendly and sustainable way of reducing damage to crops, and are an alternative to the use of chemical agents such as pesticides.[121]

In film, the idea of the predator as a dangerous if humanoid enemy is used in the 1987 science fiction horror action film Predator and its three sequels.[122][123]

See alsoEdit

NotesEdit

  1. ^ The fact that "the prey risks its life, the predator only its meal" makes the selection pressure higher on the prey than on the predator.[11]

ReferencesEdit

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  9. ^ Nilsson, Sven G.; Björkman, Christer; Forslund, Pär; Höglund, Jacob (1985). "Egg predation in forest bird communities on islands and mainland". Oecologia. 66 (4): 511–515. Bibcode:1985Oecol..66..511N. doi:10.1007/BF00379342. PMID 28310791. 
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Further readingEdit

  • Barbosa, P. and I. Castellanos (eds.) (2004). Ecology of predator-prey interactions. New York: Oxford University Press. ISBN 0-19-517120-9.
  • Curio, E. (1976). The ethology of predation. Berlin; New York: Springer-Verlag. ISBN 0-387-07720-0.