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[[File:Cheetah pursuit.jpg|thumb|A cheetah pursuing a gazelle in a predation chase]]
[[File:Cheetah pursuit.jpg|thumb|A cheetah pursuing a gazelle in a predation chase]]
'''Pursuit Predation''' is a form of [[predation]] marked by a chase from predators seeking prey. The chase can be initiated either by a predator or by prey, if alerted by a predator's presence and attempting to flee before the predator gives chase. The chase ends with either the predator capturing and consuming the prey, effectively diminishing the prey's fitness, or with the prey escaping the predators hunt, thus maintaining the prey's fitness, but leaving both prey and predator at metabolic losses. Pursuit predation is typically observed in carnivorous species within the kingdom Animalia, with some iconic examples being [[cheetah]]s, [[lion]]s, and [[Gray wolf|wolves]].
'''Pursuit Predation''' is a form of [[predation]] marked by a chase from predators seeking prey. The chase can be initiated either by a predator or by prey, if alerted by a predator's presence and attempting to flee before the predator gives chase. The chase ends with either the predator capturing and consuming the prey, effectively diminishing the prey's fitness, or with the prey escaping the predators hunt, thus maintaining the prey's fitness, but leaving both prey and predator at metabolic losses. Pursuit predation is typically observed in carnivorous species within the kingdom Animalia, with some iconic examples being the chases of [[cheetah]]s, [[lion]]s, and [[Gray wolf|wolves]].


Pursuit predation is a contrast specifically with [[ambush predation]]. While pursuit predators use a detection and pursuit phase in order to obtain prey, ambush predators use stealth to capture prey. Strength and speed are important to pursuit predators, whereas ambush predators ignore this in favor of surprise from a usually concealed location. While the two patterns of predation are not mutually exclusive, morphological differences in body plan can create a bias in an organism towards each type of predation.
Pursuit predation is an alternate predation strategy to [[ambush predation]]. While pursuit predators use a detection and pursuit phase in order to obtain prey, ambush predators use stealth to capture prey. Strength and speed are important to pursuit predators, whereas ambush predators ignore this in favor of surprise from a usually concealed location. While the two patterns of predation are not mutually exclusive, morphological differences in body plan can create a bias in an organism towards each type of predation.


== Strategy ==
== Strategy ==
There is still uncertainty as whether predators behave with a general tactic or strategy while preying.<ref>Combes SA, Salcedo MK, Pandit MM, Iwasaki JM. Capture Success and Efficiency of Dragonflies Pursuing Different Types of Prey. Integrative and Comparative Biology. 2013;56(5):787-798.</ref> Often, predators will scout potential prey, assessing prey quantity and prey density, prior to engaging in a pursuit. Certain predators pursue prey primarily among a group of conspecifics; such animals are known as [[pack hunter]](s). Other species go about the hunt alone. These two behaviors are typically due to differences in hunting success, where some groups are very successful in packs and others not as much. Predators can choose to either exhaust their metabolic resources rapidly <ref>Taylor CR , Rowntree VJ . Temperature regulation and heat balance in running cheetahs: A strategy for sprinters? Am J Physiol 224: 848-851, 1973.</ref> or pace themselves during a chase.<ref>Creel, S., & Creel, N. M. (1995, March 14). Communal hunting and pack size in African wild dogs, Lycaon pictus [Electronic version]. Animal Behavior, 50(5), 1325-1339. doi:10.1016/0003-3472(95)80048-4</ref> This choice can vary given different species of prey, different environments, and different temporal settings. Predators that rapidly exhaust their metabolic resources during a chase tend to stalk their prey, slowly moving up on their prey and trying their best not to alert it. When the predator is at closer distance (a distance that would lead easier prey capture), the predator finally gives chase.<ref>Mech, D. (1970). The Wolf: The Ecology and Behaviour of an Endangered Species. Minneapolis, MN: University of Minnesota Press.</ref> Pacing pursuit is more commonly seen in group pursuit, as individual animals do not need to exert as much energy to capture prey. However, this requires group coordination in the hunt, which may have varying degrees of success. Since groups can engage in longer chases, they often focus on separating a weaker or slower prey item during pursuit.<ref>Kruuk, H. (1972) ''The Spotted Hyena''. University of Chicago Press, Chicago IL''''</ref> Morphologically speaking, while ambush predation requires stealth,<ref>DeVries, M.S., Murphy, E.A.K. and Patek S.N. (2012). "Strike mechanics of an ambush predator: the spearing mantis shrimp". Journal of Experimental Biology 215: 4374–4384</ref> pursuit predation requires speed; pursuit predators are proportionally long-limbed and equipped with cursorial adaptations.<ref name="Janis" /> Current theories suggest that this proportionally long-limbed approach to body plan was an evolutionary countermeasure to prey adaptation.
There is still uncertainty as to whether predators behave with a general tactic or strategy while preying.<ref>Combes SA, Salcedo MK, Pandit MM, Iwasaki JM. Capture Success and Efficiency of Dragonflies Pursuing Different Types of Prey. Integrative and Comparative Biology. 2013;56(5):787-798.</ref> Often, predators will scout potential prey, assessing prey quantity and prey density, prior to engaging in a pursuit. Certain predators pursue prey primarily among a group of conspecifics; such animals are known as [[pack hunter]](s). Other species go about the hunt alone. These two behaviors are typically due to differences in hunting success, where some groups are very successful in packs and others not as much. Predators can choose to either exhaust their metabolic resources rapidly <ref>Taylor CR , Rowntree VJ . Temperature regulation and heat balance in running cheetahs: A strategy for sprinters? Am J Physiol 224: 848-851, 1973.</ref> or pace themselves during a chase.<ref>Creel, S., & Creel, N. M. (1995, March 14). Communal hunting and pack size in African wild dogs, Lycaon pictus [Electronic version]. Animal Behavior, 50(5), 1325-1339. doi:10.1016/0003-3472(95)80048-4</ref> This choice can vary given different species of prey, different environments, and different temporal settings. Predators that rapidly exhaust their metabolic resources during a chase tend to stalk their prey, slowly moving up on their prey and trying their best not to alert it. When the predator is at closer distance (a distance that would lead easier prey capture), the predator finally gives chase.<ref>Mech, D. (1970). The Wolf: The Ecology and Behaviour of an Endangered Species. Minneapolis, MN: University of Minnesota Press.</ref> Pacing pursuit is more commonly seen in group pursuit, as individual animals do not need to exert as much energy to capture prey. However, this requires group coordination in the hunt, which may have varying degrees of success. Since groups can engage in longer chases, they often focus on separating a weaker or slower prey item during pursuit.<ref>Kruuk, H. (1972) ''The Spotted Hyena''. University of Chicago Press, Chicago IL''''</ref> Morphologically speaking, while ambush predation requires stealth,<ref>DeVries, M.S., Murphy, E.A.K. and Patek S.N. (2012). "Strike mechanics of an ambush predator: the spearing mantis shrimp". Journal of Experimental Biology 215: 4374–4384</ref> pursuit predation requires speed; pursuit predators are proportionally long-limbed and equipped with cursorial adaptations.<ref name="Janis" /> Current theories suggest that this proportionally long-limbed approach to body plan was an evolutionary countermeasure to prey adaptation.


==Group Pursuers==
==Group Pursuers==


===Vertebrates===
===Vertebrates===
Group pursuers hunt with a collection of con-specifics. Group pursuit is usually seen in species of relatively high [[sociality]]. Within groups, vertebrates often seem to have defined roles in pursuit. [[African wild dog]] packs have been known to split into several smaller groups; one group initiates the chase, while the other travels ahead of the path of the prey. The group of chase initiators coordinate their chase to lead the prey towards the location of the further group where the prey's escape path will be effectively cut off.<ref>Natural World Safaris. (2015, October 16). In 10 interesting facts about African wild dogs.</ref> Pods of [[dolphins]] have been shown exhibiting a similar behavior of splitting into smaller groups. One group of the pod give chase to the fish while the other group of dolphins approach the fish from the opposite direction. This 2 pronged attack leaves the fish with only the option of jumping out of the water to escape the dolphins. However, they are left completely vulnerable in air; it is at this point when the dolphin leap out and catch the fish.<ref>REF need</ref> In lion pack hunting, each member of the hunting group is assigned a position, from left wing to right wing, in order to better obtain prey.<ref>Stander, P.E. (1992) Cooperative hunting in lions: the role of the individual. ''Behavioral Ecology and Sociobiology''. '''29''', 445-454</ref> Such specializations in role within the group are thought to occur to increase sophistication in technique; lion wing members are faster, and will drive prey toward the center where the larger, stronger, killing members of the pride will take down the prey. Many observations of group pursuers note an optimal hunting size in which certain currencies (mass of prey killed or number of prey killed) are maximized with respect to costs (kilometers chased or injuries sustained).<ref>Holekamp, K. E., Smale, L., Berg, R., & Cooper, S. M. (1996, May 22). Hunting rates and hunting success in the spotted hyena (Crocuta crocuta) [Electronic version]. Journal of Zoology, 242(1), 1-15. doi:10.1111/j.1469-7998.1997.tb02925.x</ref><ref>MacNulty, D. R., Smith, D. W., Mech, L. D., Vucetich, J. A., & Packer, C. (2011, September 29). Nonlinear effects of group size on the success of wolves hunting elk. Behavioral Ecology, 23(1), 75-82. doi:10.1093/beheco/arr159</ref> Groups size is often dependent on aspects of the environment: number of prey, number of competitors, seasonal changes, etc.
Group pursuers hunt with a collection of conspecifics. Group pursuit is usually seen in species of relatively high [[sociality]], in vertebrates individuals often seem to have defined roles in pursuit. [[African wild dog]] packs have been known to split into several smaller groups; one group initiates the chase, while the other travels ahead of the path of the prey. The group of chase initiators coordinate their chase to lead the prey towards the location of the further group where the prey's escape path will be effectively cut off.<ref>Natural World Safaris. (2015, October 16). In 10 interesting facts about African wild dogs.</ref> Pods of [[dolphins]] have been shown exhibiting a similar behavior of splitting into smaller groups. One group of the pod give chase to the fish while the other group of dolphins approach the fish from the opposite direction. This 2 pronged attack leaves the fish with only the option of jumping out of the water to escape the dolphins. However, they are left completely vulnerable in air; it is at this point when the dolphin leap out and catch the fish.<ref>REF need</ref> In lion pack hunting, each member of the hunting group is assigned a position, from left wing to right wing, in order to better obtain prey.<ref>Stander, P.E. (1992) Cooperative hunting in lions: the role of the individual. ''Behavioral Ecology and Sociobiology''. '''29''', 445-454</ref> Such specializations in role within the group are thought to occur to increase sophistication in technique; lion wing members are faster, and will drive prey toward the center where the larger, stronger, killing members of the pride will take down the prey. Many observations of group pursuers note an optimal hunting size in which certain currencies (mass of prey killed or number of prey killed) are maximized with respect to costs (kilometers chased or injuries sustained).<ref>Holekamp, K. E., Smale, L., Berg, R., & Cooper, S. M. (1996, May 22). Hunting rates and hunting success in the spotted hyena (Crocuta crocuta) [Electronic version]. Journal of Zoology, 242(1), 1-15. doi:10.1111/j.1469-7998.1997.tb02925.x</ref><ref>MacNulty, D. R., Smith, D. W., Mech, L. D., Vucetich, J. A., & Packer, C. (2011, September 29). Nonlinear effects of group size on the success of wolves hunting elk. Behavioral Ecology, 23(1), 75-82. doi:10.1093/beheco/arr159</ref> Groups size is often dependent on aspects of the environment: number of prey, number of competitors, seasonal changes, etc.


===Invertebrates===
===Invertebrates===
Matabele ants (''[[Megaponera|Megaponera analis]]''[[Megaponera|)]] raiding termite mounds
Matabele ants (''[[Megaponera|Megaponera analis]]''[[Megaponera|)]] raiding termite mounds. The predatory behavior of the bigheaded ant (''[[Pheidole megacephala]]'').
The predatory behavior of the bigheaded ant (''[[Pheidole megacephala]]'')


==Individual Pursuers==
==Individual Pursuers==
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Chicago</ref> Pursuit predation can then be supported as an adaptive mechanism for not just individual feeding success but also [[metapopulation]] persistence.
Chicago</ref> Pursuit predation can then be supported as an adaptive mechanism for not just individual feeding success but also [[metapopulation]] persistence.


==Antipredatory Adaptation to Pursuit Predation==
==Anti-predator Adaptation to Pursuit Predation==


The Evolutionary Arms Race as a vehicle for predator adaptations to anti-predator adaptations behavior is observed in flush pursuers, such as the [[painted redstart]] (''Myioborus pictus''). In such interactions, behaviors that help prey avoid more common predators will be selected for more heavily than behaviors that allow prey to avoid rarer predators. Thus, redstarts have adapted to this anti-predator adaptation.
The Evolutionary Arms Race as a vehicle for predator adaptations to anti-predator adaptations behavior is observed in flush pursuers, such as the [[painted redstart]] (''Myioborus pictus''). In such interactions, behaviors that help prey avoid more common predators will be selected for more heavily than behaviors that allow prey to avoid rarer predators. Thus, redstarts have adapted to this anti-predator adaptation.


=== The Confusion Effect and Prey Oddity ===
=== The Confusion Effect and Prey Oddity ===
In many species that fall prey to pursuit predation, gregariousness on a massive scale has evolved as a protective behavior and can be conspecific or heterospecific within a given group. This is primarily due to the Confusion Effect, which states that if prey animals congregate in large groups, predators will have more difficulty identifying and tracking specific individuals.<ref name=":0">{{Cite journal|url = |title = Oddity And the ‘Confusion Effect’ in Predation.|last = Landeau|first = Laurie|date = 1986|journal = Animal Behaviour|doi = |pmid = |access-date = |last2 = Terborgh|first2 = John}}</ref> This effect has greater influence when individuals are visually similar and less distinguishable. In groups where individuals are visually similar, there is a negative correlation between group size and predator success rates. This may mean that the overall number of attacks decreases with larger group size, or that the number of attacks per kill increases with larger group size.<ref name=":1">{{Cite journal|url = |title = Groups Confuse Predators by Exploiting Perceptual Bottlenecks: a Connectionist Model of the Confusion Effect.|last = Krakauer|first = David C.|date = 1995|journal = Behavioral Ecology and Sociobiology|doi = |pmid = |access-date = }}</ref> This is especially true in open habitats, such as [[grassland]]s or [[Pelagic zone|open ocean]] ecosystems, where view of the prey group is unobstructed, in contrast to a forest or [[reef]]. Prey species in these open environments tend to be especially gregarious, with notable examples being [[starling]]s and [[sardine]]s. When individuals of the herd are visually dissimilar, however, the success rate of predators increases dramatically.<ref name=":1" /> In one study, [[wildebeest]] on the African Savannah were selected at random and had their horns painted white. This introduced a distinction, or oddity, into the population, and researchers found that wildebeest who had white horns were preyed upon at substantially higher rates.<ref>{{Cite journal|url = |title = The Spotted Hyena. A Study Of Predation and Social Behavior.|last = Kruuk|first = Hans|date = 1973|journal = The Journal of Animal Ecology|doi = |pmid = |access-date = }}</ref> By standing out, individuals are not as easily lost in the crowd, and so predators are able to track and pursue them normally. This has been proposed as the reason why many schooling fish show little to no [[sexual dimorphism]], and why many species in heterospecific schools bear a close resemblance to other species in their school.<ref name=":0" />
In many species that fall prey to pursuit predation, gregariousness on a massive scale has evolved as a protective behavior and can be conspecific or heterospecific within a given group. This is primarily due to the [[Anti-predator adaptation|Confusion Effect]], which states that if prey animals congregate in large groups, predators will have more difficulty identifying and tracking specific individuals.<ref name=":0">{{Cite journal|url = |title = Oddity And the ‘Confusion Effect’ in Predation.|last = Landeau|first = Laurie|date = 1986|journal = Animal Behaviour|doi = |pmid = |access-date = |last2 = Terborgh|first2 = John}}</ref> This effect has greater influence when individuals are visually similar and less distinguishable. In groups where individuals are visually similar, there is a negative correlation between group size and predator success rates. This may mean that the overall number of attacks decreases with larger group size, or that the number of attacks per kill increases with larger group size.<ref name=":1">{{Cite journal|url = |title = Groups Confuse Predators by Exploiting Perceptual Bottlenecks: a Connectionist Model of the Confusion Effect.|last = Krakauer|first = David C.|date = 1995|journal = Behavioral Ecology and Sociobiology|doi = |pmid = |access-date = }}</ref> This is especially true in open habitats, such as [[grassland]]s or [[Pelagic zone|open ocean]] ecosystems, where view of the prey group is unobstructed, in contrast to a forest or [[reef]]. Prey species in these open environments tend to be especially gregarious, with notable examples being [[starling]]s and [[sardine]]s. When individuals of the herd are visually dissimilar, however, the success rate of predators increases dramatically.<ref name=":1" /> In one study, [[wildebeest]] on the African Savannah were selected at random and had their horns painted white. This introduced a distinction, or oddity, into the population, and researchers found that wildebeest who had white horns were preyed upon at substantially higher rates.<ref>{{Cite journal|url = |title = The Spotted Hyena. A Study Of Predation and Social Behavior.|last = Kruuk|first = Hans|date = 1973|journal = The Journal of Animal Ecology|doi = |pmid = |access-date = }}</ref> By standing out, individuals are not as easily lost in the crowd, and so predators are able to track and pursue them normally. This has been proposed as the reason why many schooling fish show little to no [[sexual dimorphism]], and why many species in heterospecific schools bear a close resemblance to other species in their school.<ref name=":0" />


==References==
==References==

Revision as of 18:20, 5 November 2015

File:Cheetah pursuit.jpg
A cheetah pursuing a gazelle in a predation chase

Pursuit Predation is a form of predation marked by a chase from predators seeking prey. The chase can be initiated either by a predator or by prey, if alerted by a predator's presence and attempting to flee before the predator gives chase. The chase ends with either the predator capturing and consuming the prey, effectively diminishing the prey's fitness, or with the prey escaping the predators hunt, thus maintaining the prey's fitness, but leaving both prey and predator at metabolic losses. Pursuit predation is typically observed in carnivorous species within the kingdom Animalia, with some iconic examples being the chases of cheetahs, lions, and wolves.

Pursuit predation is an alternate predation strategy to ambush predation. While pursuit predators use a detection and pursuit phase in order to obtain prey, ambush predators use stealth to capture prey. Strength and speed are important to pursuit predators, whereas ambush predators ignore this in favor of surprise from a usually concealed location. While the two patterns of predation are not mutually exclusive, morphological differences in body plan can create a bias in an organism towards each type of predation.

Strategy

There is still uncertainty as to whether predators behave with a general tactic or strategy while preying.[1] Often, predators will scout potential prey, assessing prey quantity and prey density, prior to engaging in a pursuit. Certain predators pursue prey primarily among a group of conspecifics; such animals are known as pack hunter(s). Other species go about the hunt alone. These two behaviors are typically due to differences in hunting success, where some groups are very successful in packs and others not as much. Predators can choose to either exhaust their metabolic resources rapidly [2] or pace themselves during a chase.[3] This choice can vary given different species of prey, different environments, and different temporal settings. Predators that rapidly exhaust their metabolic resources during a chase tend to stalk their prey, slowly moving up on their prey and trying their best not to alert it. When the predator is at closer distance (a distance that would lead easier prey capture), the predator finally gives chase.[4] Pacing pursuit is more commonly seen in group pursuit, as individual animals do not need to exert as much energy to capture prey. However, this requires group coordination in the hunt, which may have varying degrees of success. Since groups can engage in longer chases, they often focus on separating a weaker or slower prey item during pursuit.[5] Morphologically speaking, while ambush predation requires stealth,[6] pursuit predation requires speed; pursuit predators are proportionally long-limbed and equipped with cursorial adaptations.[7] Current theories suggest that this proportionally long-limbed approach to body plan was an evolutionary countermeasure to prey adaptation.

Group Pursuers

Vertebrates

Group pursuers hunt with a collection of conspecifics. Group pursuit is usually seen in species of relatively high sociality, in vertebrates individuals often seem to have defined roles in pursuit. African wild dog packs have been known to split into several smaller groups; one group initiates the chase, while the other travels ahead of the path of the prey. The group of chase initiators coordinate their chase to lead the prey towards the location of the further group where the prey's escape path will be effectively cut off.[8] Pods of dolphins have been shown exhibiting a similar behavior of splitting into smaller groups. One group of the pod give chase to the fish while the other group of dolphins approach the fish from the opposite direction. This 2 pronged attack leaves the fish with only the option of jumping out of the water to escape the dolphins. However, they are left completely vulnerable in air; it is at this point when the dolphin leap out and catch the fish.[9] In lion pack hunting, each member of the hunting group is assigned a position, from left wing to right wing, in order to better obtain prey.[10] Such specializations in role within the group are thought to occur to increase sophistication in technique; lion wing members are faster, and will drive prey toward the center where the larger, stronger, killing members of the pride will take down the prey. Many observations of group pursuers note an optimal hunting size in which certain currencies (mass of prey killed or number of prey killed) are maximized with respect to costs (kilometers chased or injuries sustained).[11][12] Groups size is often dependent on aspects of the environment: number of prey, number of competitors, seasonal changes, etc.

Invertebrates

Matabele ants (Megaponera analis) raiding termite mounds. The predatory behavior of the bigheaded ant (Pheidole megacephala).

Individual Pursuers

Vertebrates

Big cat species are prevalent individual pursuers. Cheetahs are widely known as the fastest terrestrial animal, with measures of speeds from 68-75 mph.[13] Their speed and acceleration give them a great advantage in pursuit. However, their speed can only be sustained for short periods of time. Overheating can become a fatal consequence of prolonged fast paced chase by cheetahs. As such, cheetahs, like many other big cats, will stalk their prey until they can get as close as possible without being detected before finally giving chase.

Flush pursuers are avian pursuers that purposefully alert prey of their presence to enhance prey capture. The Painted redstart (Myioborus pictus) is one of the most well documented flush pursuers. When flies, prey for redstarts, are alerted of the presence of predators, they respond by fleeing. Redstarts take advantage of this antipredator response by spreading and orienting their easily noticeable wings and tails, alerting the flies, but only when they are in a position where the flies' escape path intersects with the redstart's central field of vision. Once the flies begin to escape, the redstart begins to chase. This strategy greatly increases the number of prey captured. It has been proposed possible that redstarts exploit two aspects of the visual sensitivity of prey: sensitivity to the location of the stimulus in the prey's visual field and sensitivity to the direction of stimulus environment.[14] The effectiveness of this pursuit can also be explained by "rare enemy effect", an evolutionary consequence of multi-species predator-prey interactions.[14]

Invertebrates

Dragonflies are skilled aerial pursuers; they have a 97% success prey capture rate.[15] This success rate is a consequence of the "decision-making" of which prey to pursue based on initial conditions. Observations of several species of perching dragonflies show more pursuit initiations at larger starting distances for larger size prey species than for much smaller prey. Further evidence points to a potential foraging strategy of dragonflies choosing to pursue larger prey at any given opportunity, due to more substantial metabolic rewards. This is in spite of the fact that larger prey typically stipulate faster prey and less successful pursuits. Dragonflies high success prey capture rate may also be due to their "interception" foraging method, unlike the tracking foraging methods, in which predators focus on closing in on the current position of their prey. Instead the interception method has the dragonfly seeking the position directly ahead of their prey as a way of surmising a prey's future location. Perching dragonflies (the Libellulidae family), the largest family of dragonflies, have been observed "staking out" high density prey spots prior to pursuit.[16] There are no noticeable distinctions in prey capture efficiency between male and female dragonflies. Further, percher dragonflies are more likely to engage in pursuit when prey come within a subtend angle of around 1-2 degrees. Angles greater than this are outside of a dragonflies visual range.

Evolutionary Basis of the Behavior

Evolution as a countermeasure

Current theory on the evolution of pursuit predation suggests that the behavior is actually an evolutionary countermeasure to prey adaptation. Prey animals vary in their likelihood to avoid predation, and it is predation failure that drives evolution of both prey and predator.[17] Predation failure rates vary wildly across the animal kingdom; raptorial birds can fail anywhere from 20% to 80% of the time in predation, while predatory mammals usually fail more than half the time.[17] Prey adaptation drives these low rates in three phases: the detection phase, the pursuit phase, and the resistance phase.[18] The pursuit phase drove the evolution of distinct behaviors for pursuit predation.

Phases of Pursuit Predation

As selective pressure on prey is higher than on predators [17] adaptation usually occurs in prey long before the reciprocal adaptations in predators. Evidence in the fossil record supports this, with no evidence of modern pursuit predators until the late Tertiary.[7] Certain adaptations, like long limbs in ungulates, that were thought to be adaptive for speed against predatory behavior have been found to predate predatory animals by over 20 million years. Because of this, modern pursuit predation is an adaptation that may have evolved separately and much later as a need for more energy in colder and more arid climates.[7] Longer limbs in predators, the key morphological adaptation required for lengthy pursuit of prey, is tied in the fossil record to the late Tertiary. It is now believed that modern pursuit predators like the wolf and lion evolved this behavior around this time period as a response to ungulates increasing feeding range.[7] As ungulate prey moved into a wider feeding range to discover food in response to changing climate, predators evolved the longer limbs and behavior necessary to pursue prey across larger ranges. In this respect, pursuit predation is not co-evolutionary with prey adaptation, but a direct response to prey. Prey adaptation to climate is the key formative reason for evolving the behavior and morphological necessities of pursuit predation.

In addition to serving as a countermeasure to prey adaptation, pursuit predation has evolved in some species as an alternative, facultative mechanism for foraging. For example, polar bears typically act as specialized predators of seal pups and operate in a manner closely predicted by the optimal foraging theory. However, they have been seen to occasionally employ more energy-inefficient pursuit predation tactics on flightless geese. This alternative predatory strategy may serve as a back-up resource when optimal foraging is circumstantially impossible, or may even be a function of filling dietary needs.[19]

Evolution from an Ecological Basis

Pursuit predation revolves around a distinct movement interaction between predator and prey; as prey move to find new foraging areas, predators should move with them. Predators congregate in areas of high prey density,[20] and prey should then avoid these areas in turn.[21] Because of these interactions, Spatial patterns of predators and prey are important in preserving population size. Prey attempts to avoid predation and find food are coupled with predator attempts to find food and compete with other predators. These interactions act to preserve populations.[22] Models of spatial patterns and synchrony of predator-prey relationships can be used as support for the evolution of pursuit predation as one mechanism to preserve these population mechanics. By pursuing prey over long distances, predators actually improve longterm survival of both their own population and prey population through population synchrony. Pursuit predation acts to even out population fluctuations by moving predatory animals from areas of high predator density to low predator density, and low prey density to high prey density. This keeps migratory populations in synchrony, which increases metapopulation persistence.[22] Pursuit predation’s effect on population persistence is more marked over larger travel ranges. Predator and prey levels are usually more synchronous in predation over larger ranges, as population densities have more ability to even out.[23] Pursuit predation can then be supported as an adaptive mechanism for not just individual feeding success but also metapopulation persistence.

Anti-predator Adaptation to Pursuit Predation

The Evolutionary Arms Race as a vehicle for predator adaptations to anti-predator adaptations behavior is observed in flush pursuers, such as the painted redstart (Myioborus pictus). In such interactions, behaviors that help prey avoid more common predators will be selected for more heavily than behaviors that allow prey to avoid rarer predators. Thus, redstarts have adapted to this anti-predator adaptation.

The Confusion Effect and Prey Oddity

In many species that fall prey to pursuit predation, gregariousness on a massive scale has evolved as a protective behavior and can be conspecific or heterospecific within a given group. This is primarily due to the Confusion Effect, which states that if prey animals congregate in large groups, predators will have more difficulty identifying and tracking specific individuals.[24] This effect has greater influence when individuals are visually similar and less distinguishable. In groups where individuals are visually similar, there is a negative correlation between group size and predator success rates. This may mean that the overall number of attacks decreases with larger group size, or that the number of attacks per kill increases with larger group size.[25] This is especially true in open habitats, such as grasslands or open ocean ecosystems, where view of the prey group is unobstructed, in contrast to a forest or reef. Prey species in these open environments tend to be especially gregarious, with notable examples being starlings and sardines. When individuals of the herd are visually dissimilar, however, the success rate of predators increases dramatically.[25] In one study, wildebeest on the African Savannah were selected at random and had their horns painted white. This introduced a distinction, or oddity, into the population, and researchers found that wildebeest who had white horns were preyed upon at substantially higher rates.[26] By standing out, individuals are not as easily lost in the crowd, and so predators are able to track and pursue them normally. This has been proposed as the reason why many schooling fish show little to no sexual dimorphism, and why many species in heterospecific schools bear a close resemblance to other species in their school.[24]

References

  1. ^ Combes SA, Salcedo MK, Pandit MM, Iwasaki JM. Capture Success and Efficiency of Dragonflies Pursuing Different Types of Prey. Integrative and Comparative Biology. 2013;56(5):787-798.
  2. ^ Taylor CR , Rowntree VJ . Temperature regulation and heat balance in running cheetahs: A strategy for sprinters? Am J Physiol 224: 848-851, 1973.
  3. ^ Creel, S., & Creel, N. M. (1995, March 14). Communal hunting and pack size in African wild dogs, Lycaon pictus [Electronic version]. Animal Behavior, 50(5), 1325-1339. doi:10.1016/0003-3472(95)80048-4
  4. ^ Mech, D. (1970). The Wolf: The Ecology and Behaviour of an Endangered Species. Minneapolis, MN: University of Minnesota Press.
  5. ^ Kruuk, H. (1972) The Spotted Hyena. University of Chicago Press, Chicago IL'
  6. ^ DeVries, M.S., Murphy, E.A.K. and Patek S.N. (2012). "Strike mechanics of an ambush predator: the spearing mantis shrimp". Journal of Experimental Biology 215: 4374–4384
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  13. ^ Sharp, N. C. (2009, March 24). Timed running speed of a cheetah. Journal of Zoology, 241(3), 493-494. doi:10.1111/j.1469-7998.1997.tb04840.x
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  15. ^ Olberg RM, Worthington AH, Venator KR J Comp Physiol A. 2000 Feb; 186(2):155-62.
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  19. ^ Iles, D. T., Peterson, S. L., Gormezano, L. J., Koons, D. N., & Rockwell, R. F. (2013). Terrestrial predation by polar bears: Not just a wild goose chase. Polar Biology, 36(9), 1373-1379. doi:http://dx.doi.org/10.1007/s00300-013-1341-5
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  22. ^ a b Li, Z. Z., Gao, M., Hui, C., Han, X. Z., & Shi, H. (2005). Impact of predator pursuit and prey evasion on synchrony and spatial patterns in metapopulation. Ecological Modelling, 185(2), 245-254.
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