Predation
thumb|upright=1.35|A polar bear as the predator feeding on a bearded seal
thumb|upright=1.35|Meat ants cooperating to feed on a cicada far larger than themselves

In an ecosystem, predation is a biological interaction where a predator (an organism that is hunting) feeds on its prey (the organism that is attacked).Begon, M., Townsend, C., Harper, J. (1996). Ecology: Individuals, populations and communities (Third edition). Blackwell Science, London. , , . Predators may or may not kill their prey prior to feeding on it, but the act of predation often results in the death of the prey and the eventual absorption of the prey's tissue through digestion. Predation is often, though not always, carnivory, one of several heterotrophic consumer-resource interactions. Different feeding behaviors such as parasitism, parasitoidism, micropredation and predation form a continuum rather than being entirely separate strategies. Predation strategies can be classified by trophic level or diet, by specialization, and by the predator's interaction with prey.

Coevolutionary selective pressures imposed on each other by predator and prey often result in an evolutionary arms race, resulting in antipredator adaptations in the prey and adaptations that improve hunting efficiency in the predator.

 By function 

Predators can be classified by their consumer-resource interactions with their prey. Two factors are considered here: how close the predator and prey (or host) are, and whether the prey is directly killed by the predator, where true predation and parasitoidism involve certain death.

 True predation 
thumb|Humpback whales filtering thousands of krill from seawater, using their baleen systems

A true predator is one that kills and eats another living thing. Predators may hunt actively for prey in pursuit predation, or sit and wait for prey to approach within striking distance in ambush predation. Some predators kill large prey and dismember or chew it prior to eating it, as do humans; others may eat their prey whole, as do bottlenose dolphins swallowing fish, white storks swallowing frogs, or baleen whales swallowing thousands of krill or small fish at once. Some predators use venom to subdue their prey before the predator ingests it, as in the box jellyfish, while the venom of rattlesnakes and some spiders also helps to digest the prey. Seed and egg predation are true predation, as seeds and eggs are potential organisms.

 Grazing 

Grazing animals generally do not kill their prey, but like predators, they live by feeding on other organisms. While some herbivores like zooplankton live on unicellular phytoplankton and therefore inevitably kill what they eat, in a relationship sometimes called predation, many others including cattle and sheep only eat a part of the plants that they graze. Many species of plant are adapted to regrow after grazing damage. For example, the growing meristems of grasses are not at the tips as they are in most flowering plants, but at the base of the leaves. Similarly, kelp is grazed in subtidal kelp forests, but continuously regrows from a meristem at the base of the blade where it joins the stipe. Animals may also be 'grazed' upon; micropredators such as female mosquitoes land on hosts briefly to feed on blood. Herbivore-plant interactions, as with predator-prey interactions, have driven plants to evolve defences such as thorns and chemicals to dissuade grazing.

 Parasitism 

thumb|Tapeworm Taenia solium, a parasite, has hooks and suckers on its head to attach to its host. 

Parasites, like predators, live by feeding on another organism. The entomologist E. O. Wilson has characterised parasites as "predators that eat prey in units of less than one". Parasites often do not kill their hosts, the exception being parasitoids which always do, thus blurring the line between parasitism and predation. Equally, small parasites such as mosquitoes exploit their hosts much as micropredators such as moth caterpillars on an oak tree and grazers do, though endoparasites in particular have a close association with their host species; again there is essentially a continuum between these feeding interactions.

 Parasitoidism 

Parasitoids are organisms living in or on their host and feeding directly upon it, eventually leading to its death. They are much like parasites in their close associations with their hosts. Unlike typical parasites, they always kill their hosts, but often not instantly. Parasitoid wasps are solitary insects that live a free life as adults, laying eggs on or in other insects such as lepidopteran caterpillars. The wasp larvae feed on the growing host, eventually killing it. Parasitoids make up as much as 10% of all insect species.

Ecological interactions
Specialization
thumb|Platydemus manokwari is a flatworm predator of land snails.

Predators are often highly specialized, where for example the Eurasian lynx only hunts small ungulates. Others such as leopards are more opportunistic generalists. 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 to be more specialized as the prey are more conspicuous and can be found more quickly.

 Trophic level 
thumb|A secondary consumer in action: 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. For example, a lion, an apex predator (at the top of its food chain) 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. 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.

 As competition 
An alternative view offered by Richard Dawkins is of predation as a form of competition: the genes of both the predator and prey are competing for the body (or 'survival machine') of the prey organism. This is best understood in the context of the gene centered view of evolution. Another manner in which predation and competition are connected is throughout intraguild predation. Intraguild predators are those that kill and eat other predators of different species at the same trophic level, and thus that are potential competitors.

 Biodiversity 
thumb|upright=1.3|Riparian willow recovery at Blacktail Creek, Yellowstone National Park, after reintroduction of wolves

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.Changing the distribution of predators and prey in an ecosystem can turn things upside down. March 1, 2013 Scientific American  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.Botkin, D. and E. Keller (2003). Environmental Science: Earth as a living planet. John Wiley & Sons. . P.2.

The elimination of wolves from Yellowstone National Park had profound impacts on the trophic pyramid. 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 in riparian areas, which protected beavers from having their food sources encroached upon. The removal of wolves had a direct effect on beaver populations, as their habitat became territory for grazing.William J. Ripple and Robert L. Beschta. "Wolves and the Ecology of Fear: Can Predation Risk Structure Ecosystems?" 2004. Furthermore, predation keeps hydrological features such as creeks and streams in normal working order. Increased browsing on willows and conifers along Blacktail Creek due to a lack of predation caused channel incision because the beavers helped slow the water down, allowing soil to stay in place.

 Optimal foraging 

While successful predation results in a gain of energy, hunting invariably involves energetic costs. When not hungry, predators stop hunting, since the costs outweigh the benefits. For instance, a large well-fed predatory fish like a shark in an aquarium ignores prey fish nearby. Surplus killing represents a deviation from this  behaviour. The treatment of consumption in terms of cost-benefit analysis is called optimal foraging theory, and has been successful in explaining much animal behavior.

Social predation
thumb|upright=1.6|Wolves cooperating to hunt bison

In social predation, a group of predators cooperates to kill creatures larger than those they could overpower singly. Social predators such as ions, hyenas, and wolves collaborate to catch and kill large herbivores. By hunting socially chimpanzees can catch colobus monkeys that would readily escape an individual hunter, while a cooperating group of Harris hawks can trap rabbits.

Size-selective predation

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.

 Antipredator adaptations 

thumb|upright|left|Springbok stotting to advertise its ability to escape

Many antipredator adaptations have evolved in prey populations due to the selective pressures of predation over long periods of time. Some species mob predators cooperatively. Others such as Thomson's gazelle stot to  signal to predators such as cheetahs that they will have an unprofitable chase. Many prey animals are aposematically colored or patterned as a warning to predators that they are distasteful or able to defend themselves.Bowers, M. D., Irene L. Brown, and Darryl Wheye. "Bird Predation as a Selective Agent in a Butterfly Population." Evolution 39.1 (1985): 93-103. Such distastefulness or toxicity is brought about by chemical defenses, found in a wide range of prey, especially insects, but the skunk is a dramatic mammalian example. Chemical defences can include toxins, such as bitter compounds in leaves, often absorbed by leaf-eating insects such as caterpillars and used to dissuade potential predators.Toxins and venoms. Author: Edmund D.Brodie Mountain Lake Biological Station, Department of Biology, University of Virginia, Charlottesville, VA 22904-4328, USA

thumb|Camouflage of the dead leaf mantis makes it less visible to both its predators and its prey.

Camouflage makes use of coloration, shape, and pattern to misdirect the visual sensory mechanisms of predators, enabling prey to remain unrecognized.Merilaita, S., Scott-Samuel, N. E., & Cuthill, I. C. (2017). How camouflage works. Philosophical Transactions Of The Royal Society B: Biological Sciences, 372(1724), 1-9. doi:10.1098/rstb.2016.0341 Among the many mechanisms of camouflage are blending in with ones background, countershading, and disruptive coloration. 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 wing markings that resemble eyes. When a predator disturbs the insect, it reveals its hind wings, startling the predator and giving it time to escape.

 Predator camouflage and aggressive mimicry 
thumb|A camouflaged predator: snow leopard in Ladakh (Photo by Tashi Lonchay)
thumb|Striated frogfish uses camouflage and aggressive mimicry in the form of a fishing rod-like lure on its head to attract prey.

Predators use stealth, camouflage and aggressive mimicry to improve their hunting efficiency. 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. Female Photuris 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.

 Population dynamics 

Predators tend to lower the survival and fecundity of their prey, and 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 modelled using the Lotka–Volterra equations. These provide a mathematical model for the cycling of predator and prey populations. Predators tend to select young, weak, and ill individuals.

 Evolution
Predation appears to have become a major selection pressure shortly before the Cambrian 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 .

 Humans 
thumb|Humans and dogs as a predatory team

Humans are to some extent predatory, fishing, hunting and trapping animals using weapons and tools. They also use other predatory species, such as dogs, cormorants, and falcons to catch prey for food or for sport.
 
In biological pest control, predators from a pest's natural range are introduced to control populations, at the risk of causing unforeseen problems. Natural predators 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.

 See also 
 Bird of prey
  Built for the Kill, a nature series on predators
 Overpopulation in wild animals
 Predator–prey reversal
 Prey drive
 Refuge (ecology)
 Wa-Tor

 References 

 Further reading 
 Barbosa, P. and I. Castellanos (eds.) (2004). Ecology of predator-prey interactions. New York: Oxford University Press. .
 Curio, E. (1976). The ethology of predation. Berlin; New York: Springer-Verlag. .

 External links 

 Wolfram Demonstrations Project:  Predator-Prey Equations by Eric W. Weisstein
Predators, three articles by Olivia Judson, NY Times, Sept. & Oct., 2009

Category:Predation
Category:Biological pest control