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'''Pursuit Predation''' is a form of [[predation]] marked by a chase from predators seeking prey. The chase can be initiated either by predators or by prey, alerted by a predators presence, that attempt to flee before predators give chase. The chase ends with either the predator's capture and consumption of 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 is typically observed in carnivorous organisms, with animals being the most transparent examples of pursuit predators. |
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'''Communal roosting''' is practiced by birds, bats and some insects when large flocks or colonies roost together usually in trees with several hundred on each.<ref>Barclay, Robert MR. "Night roosting behavior of the little brown bat, Myotis lucifugus." ''Journal of Mammalogy'' 63.3 (1982): 464-474.</ref><ref>Klug, Brandon J., and Robert MR Barclay. "Thermoregulation during reproduction in the solitary, foliage-roosting hoary bat (Lasiurus cinereus)." ''Journal of Mammalogy'' 94.2 (2013): 477-487.</ref> |
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<big>'''I. Strategy '''</big> |
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Several communal roosting trees can be located within densely populated cities nowadays where common birds like [[house sparrow]]s and [[starling]]s etc. can be seen roosting in large numbers. |
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In depth analysis of predation: beginning, during, and afterwards, important to make distinctions from not only ambush, but other types if necessary. 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> Certain predators will scout potential prey, assessing prey quantity, prey density, and, if a member of predator group, the spread of other conspecfics( ). During a chase, predators may either exhaust their energy rapidly or pace themselves when anticipating a long pursuit. |
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Some benefits as a result of communal roosting are reduced predation risk, increased breading interactions and reduced cost in territory defence.<ref>Agosta, Salvatore J. "Habitat use, diet and roost selection by the big brown bat (Eptesicus fuscus) in North America: a case for conserving an abundant species." ''Mammal Review'' 32.3 (2002): 179-198.</ref> |
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<big>'''II. Evolutionary Basis of the Behavior</big> |
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== The Evolution of Communal Roosting == |
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'''A) Evolution as a countermeasure'''''' |
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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 <ref name = "Vermeij"> Geerat Vermeij,The American Naturalist |
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=== The Information Center Hypothesis (ICH) === |
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Vol. 120, No. 6 (Dec., 1982), pp. 701-720 </ref>. 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 <ref name="Vermeij" />. Prey adaptation drives these low rates in three phases: the detection phase, the pursuit phase, and the resistance phase <ref name = "Holling"> Holling, C. S. (1966). The functional response of invertebrate predators to prey density. Memoirs of the Entomological Society of Canada, 98(S48), 5-86. </ref>. The pursuit phase drove the evolution of distinct behaviors for pursuit predation. As selective pressure on prey is higher than on predators <ref name="Vermeij" /> 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 <ref name = "Janis"> Janis, C. M., & Wilhelm, P. B. (1993). Were there mammalian pursuit predators in the Tertiary? Dances with wolf avatars. Journal of Mammalian Evolution, 1(2), 103-125.</ref>. 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<ref name="Janis" />. 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 <ref name="Janis" />. 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. |
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=== The Two Strategies Hypothesis === |
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The Two Strategies Hypothesis was put forth by Patrick Weatherhead in 1983 as an alternative to the then popular Information Center Hypothesis. The Two Strategies Hypothesis proposes that instead of joining roosts due to increased foraging capabilities, different individuals within a communal roost participate in the roost for different reasons. This hypothesis explains that older more experienced foragers remain within a communal roost due to the fact that they are considered dominant, and therefore able to obtain the safest roosts, with the less dominant and unsuccessful foragers acting as a buffer to predation. This is similar to the [[selfish herd theory]], which states that individuals within herds will utilize conspecifics to avoid predation. The younger individuals will remain with the roost as they still gain some safety from predation through the dilution factor, as well as the ability to learn from the more experienced foragers.<ref>{{Cite journal|url = http://www.jstor.org.proxy.lib.duke.edu/stable/2461125?pq-origsite=summon&seq=2#page_scan_tab_contents|title = Two Principal Strategies in Avian Communal Roosts|last = Weatherhead|first = Patrick|date = February 1983|journal = The American Naturalist|doi = |pmid = |access-date = October 15, 2015|pages = pp. 237-247}}</ref> A study of roosting rooks (''[[Rook (bird)|Corvus frugilegus]]'') supports this hypothesis, showing that within rook communal roosts there exists an inherent hierarchy, with the most dominant occupying the roosts highest in the tree, and thus safer from terrestrial predators.<ref>{{Cite journal|url = http://onlinelibrary.wiley.com/doi/10.1111/j.1469-7998.1977.tb04167.x/abstract|title = The social and spatial organization of winter communal roosting in Rooks (Corvus frugilegus)|last = Swingland|first = Ian R.|date = August 1977|journal = Journal of Zoology|doi = |pmid = |access-date = October 15, 2015|pages = pp. 509-528}}</ref> |
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'''B) Evolution from an Ecological Basis''' |
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=== The Recruitment Center Hypothesis (RCH) === |
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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 <ref name = "Krebs"> Krebs, J. R. (1978). Optimal foraging: decision rules for predators. Behavioural ecology: an evolutionary approach, 23-63.</ref>, and prey should then avoid these areas in turn <ref name = "Sih"> The Behavioral Response Race Between Predator and Prey, Andrew Sih, The American Naturalist, Vol. 123, No. 1 (Jan., 1984), pp. 143-150</ref>. 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 <ref name = "Li">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. </ref>. 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 <ref name = "Li">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. </ref>. 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 <ref name = "Leggett"> Rose, G. A., & Leggett, W. C. (1990). The importance of scale to predator-prey spatial correlations: an example of Atlantic fishes. Ecology, 33-43. |
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== Potential Benefits of Communal Roosting == |
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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. |
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==Invertebrates== |
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== Potential Costs of Communal Roosting == |
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[[Dragonflies]] are skilled aerial pursuer; they have a 97% success prey capture rate. <ref> Olberg RM, Worthington AH, Venator KR J Comp Physiol A. 2000 Feb; 186(2):155-62. </ref> 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. <ref> Perching dragonflies (the [[Libellulidae]] family), the largest family of dragonflies, have been observed "staking out" high density prey spots prior to pursuit. <ref> Combes, S. A., M. K. Salcedo, M. M. Pandit, and J. M. Iwasaki. "Capture Success and Efficiency of Dragonflies Pursuing Different Types of Prey." Integrative and Comparative Biology 53.5 (2013): 787-98. Web. </ref> There are no noticeable distinctions in prey capture efficiency between male and female dragonflies. Further, percher dragonflies are 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. |
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== Examples of Communal Roosts in Extant Species == |
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=== Communal Roosting in Birds === |
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Communal roosting has been observed in numerous avian species. Acorn woodpeckers (''[[Acorn woodpecker|Melanerpes formicivorus]]'') are known to form communal roosts during the winter months, sharing their body heat and decreasing the thermoregulatory demands on all individuals in the roost.<ref>Plessis, Ma du., Morné A., Wesley W. Weathers, and Walter D. Koenig. “Energetic benefits of communal roosting by acorn woodpeckers during the nonbreeding season.” ''Condor'' (1994): 631–637.</ref> The tree swallow ([[Tree swallow|''Tachycineta bicolor'']]) is known to form communal roosts and exhibits high roost fidelity and it is believed that high conspecific attraction leads to the forming of communal roosts.<ref>Laughlin, A. J., D. R. Sheldon, D. W. Winkler, and C. M. Taylor. "Behavioral Drivers of Communal Roosting in a Songbird: A Combined Theoretical and Empirical Approach." ''Behavioral Ecology'' 25.4 (2014): 734-43. Web. 29 Sept. 2015.</ref> Red-billed choughs ([[Red-billed chough|''Pyrrhocorax pyrrhocorax'']]) roost in either a main roost or a sub roost. Main roosts are used constantly, whereas the sub roosts are used irregularly by individuals lacking both a mate and territory. These sub roosts are believed to help improve the ability of non-breeding choughs to find a mate and increase their territory.<ref>Blanco, Guillermo and Jose L. Tella. “Temporal, spatial and social segregation of red-billed choose between two types of communal roost: a role for mating and territory acquisition.” ''The Association for the Study of Animal Behaviour'' 57 (1999): 1219-1227. </ref> Interspecies roosts have also been observed in nature. Great egrets (''[[Great egret|Ardea alba]]''), little blue herons (''[[Little blue heron|Egretta caerulea]]''), tricolored herons (''[[Tricolored heron|Egretta tricolor]]''), and the snowy egret (''[[Snowy egret|Egretta thula]]'') are known to form large communal roosts in San Blas, Mexico. It has been shown that the snowy egret determines the general location of the roost due to the fact that the other three species rely on it for its abilities to find food sources. In these roosts there is often a hierarchical system, where the more dominant species (in this case the snowy egret) will typically occupy the more desirable higher perches.<ref>Burger, J., et al. "Intraspecific and interspecific interactions at a mixed species roost of ciconiiformes in San Blas, Mexico."''Biology of Behaviour''" (1977): 309-327.</ref> Interspecies roosts have also been observed among other avian species.<ref>Burger, Joanna. "A model for the evolution of mixed-species colonies of Ciconiiformes." ''Quarterly Review of Biology'' (1981): 143-167.</ref><ref>Munn, Charles A., and John W. Terborgh. "Multi-species territoriality in Neotropical foraging flocks." ''Condor'' (1979): 338-347.</ref> |
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=== Communal Roosting in Insects === |
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Communal roosting has also been well documented among insects, particularly butterflies. The passion-vine butterfly (''[[Heliconius erato|Heliconius erato)]]'' is known to form large nocturnal roosts. It is believed that these roosts deter potential predators due to the fact that predators infrequently attack large roosts.<ref>Finkbeiner, Susan D., Adriana D. Briscoe, and Robert D. Reed. “The benefit of being a social butterfly: communal roosting deters predation.” Proceedings of the Royal Society of London B: ''Biological Sciences'' 279.1739 (2012): 2769–2776.</ref> Communal roosting has also been observed in south peruvian tiger beetles of the genus [[Beetle|''Coleoptera'' and ''Cicindelidae'']]. These species of tiger beetle have been observed to form communal roosts comprising anywhere from 2-9 individuals at night and disbanding during the day. It is hypothesized that these beetles roost high in the treetops in order to avoid ground-based predators.<ref>Pearson, David L., and Joseph J. Anderson. "Perching heights and nocturnal communal roosts of some tiger beetles (Coleoptera: Cicindelidae) in southeastern Peru." ''Biotropica'' (1985): 126-129.</ref> |
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=== Communal Roosting in Mammals === |
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While there are few observations of communal roosting mammals, the trait has been seen in several species of bats. The little brown bat (''[[Little brown bat|Myotis lucifugus]]'') is known to participate in communal roosts of up to 37 during cold nights in order to decrease thermoregulatory demands, with the roost disbanding at daybreak.<ref>Barclay, Robert MR. "Night roosting behavior of the little brown bat, Myotis lucifugus." ''Journal of Mammalogy'' 63.3 (1982): 464-474.</ref> Several other species of bats, including the hoary bat (''[[Hoary bat|Lasiurus cinereu]]s'') and the big brown bat (''[[Big brown bat|Eptesicus fuscus]]'') have also been observed to roost in maternal colonies in order to reduce the thermoregulatory demands on both the lactating mothers and juveniles.<ref>Klug, Brandon J., and Robert MR Barclay. "Thermoregulation during reproduction in the solitary, foliage-roosting hoary bat (Lasiurus cinereus)." ''Journal of Mammalogy'' 94.2 (2013): 477-487.</ref><ref>Agosta, Salvatore J. "Habitat use, diet and roost selection by the big brown bat (Eptesicus fuscus) in North America: a case for conserving an abundant species." ''Mammal Review'' 32.3 (2002): 179-198.</ref> |
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== See also == |
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* [[Communal breeding]] |
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* [[Habitat]] |
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* [[Ecology]] |
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* [[Ecosystem]] |
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* [[Habitat conservation]] |
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* [[Habitat fragmentation]] |
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* [[Reproduction]] |
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* [[Mating system]] |
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* [[Cooperative breeding]] |
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*[[Heliconius charithonia]] |
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==References== |
==References== |
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<references />{{reflist}} |
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* |
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==Notes== |
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{{reflist}} |
{{reflist}} |
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{{uncat|date=October 2015}} |
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==External links== |
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[[Category:Ecology]] |
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{{ecology-stub}} |
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Revision as of 16:02, 20 October 2015
Pursuit Predation is a form of predation marked by a chase from predators seeking prey. The chase can be initiated either by predators or by prey, alerted by a predators presence, that attempt to flee before predators give chase. The chase ends with either the predator's capture and consumption of 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 is typically observed in carnivorous organisms, with animals being the most transparent examples of pursuit predators.
I. Strategy
In depth analysis of predation: beginning, during, and afterwards, important to make distinctions from not only ambush, but other types if necessary. There is still uncertainty as whether predators behave with a general tactic or strategy while preying. [1] Certain predators will scout potential prey, assessing prey quantity, prey density, and, if a member of predator group, the spread of other conspecfics( ). During a chase, predators may either exhaust their energy rapidly or pace themselves when anticipating a long pursuit.
II. Evolutionary Basis of the Behavior
A) 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 [2]. 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 [2]. Prey adaptation drives these low rates in three phases: the detection phase, the pursuit phase, and the resistance phase [3]. The pursuit phase drove the evolution of distinct behaviors for pursuit predation. As selective pressure on prey is higher than on predators [2] 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 [4]. 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[4]. 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 [4]. 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.
B) 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 [5], and prey should then avoid these areas in turn [6]. 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 [7]. 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 [7]. 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 [8]. Pursuit predation can then be supported as an adaptive mechanism for not just individual feeding success but also metapopulation persistence.
Invertebrates
Dragonflies are skilled aerial pursuer; they have a 97% success prey capture rate. [9] 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. Cite error: A <ref> tag is missing the closing </ref> (see the help page). There are no noticeable distinctions in prey capture efficiency between male and female dragonflies. Further, percher dragonflies are 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.
References
- ^ 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.
- ^ a b c Geerat Vermeij,The American Naturalist Vol. 120, No. 6 (Dec., 1982), pp. 701-720
- ^ Holling, C. S. (1966). The functional response of invertebrate predators to prey density. Memoirs of the Entomological Society of Canada, 98(S48), 5-86.
- ^ a b c Janis, C. M., & Wilhelm, P. B. (1993). Were there mammalian pursuit predators in the Tertiary? Dances with wolf avatars. Journal of Mammalian Evolution, 1(2), 103-125.
- ^ Krebs, J. R. (1978). Optimal foraging: decision rules for predators. Behavioural ecology: an evolutionary approach, 23-63.
- ^ The Behavioral Response Race Between Predator and Prey, Andrew Sih, The American Naturalist, Vol. 123, No. 1 (Jan., 1984), pp. 143-150
- ^ 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.
- ^ Rose, G. A., & Leggett, W. C. (1990). The importance of scale to predator-prey spatial correlations: an example of Atlantic fishes. Ecology, 33-43. Chicago
- ^ Olberg RM, Worthington AH, Venator KR J Comp Physiol A. 2000 Feb; 186(2):155-62.