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{{short description|Animal and human predatory behavior}}
{{short description|Animal and human predatory behavior}}
[[File:Stoat and chipmunks.png|thumb|right|A [[stoat]] surplus killing [[chipmunks]] ([[Ernest Thompson Seton]], 1909)]]
[[File:Stoat and chipmunks.png|thumb|right|A [[stoat]] surplus killing [[chipmunks]] ([[Ernest Thompson Seton]], 1909)]]
[[File:Multiple sheep kill.jpg|thumb|Multiple sheep killed by a [[Cougar|cougar]]]]
[[File:Multiple sheep kill.jpg|thumb|Multiple sheep killed by a [[cougar]]]]
'''Surplus killing''', also known as '''excessive killing''', '''killing for sport''', '''henhouse syndrome''',<ref name="LSM148" /><ref>{{cite book|last=Moskowitz|first=David|title=Wolves in the Land of Salmon.|date=4 February 2013|publisher=Timber Press|isbn=978-1604692273|pages=144}}</ref> or '''overkill''',<ref>{{Cite journal|last=Mysterud|first=Ivar|date=1980|title=Bear Management and Sheep Husbandry in Norway, with a Discussion of Predatory Behavior Significant for Evaluation of Livestock Losses|url=http://www.jstor.org/stable/3872873|journal=Bears: Their Biology and Management|volume=4|pages=233–241|doi=10.2307/3872873|jstor=3872873|issn=1936-0614}}</ref> is a common behavior exhibited by [[predator]]s, in which they kill more [[prey]] than they can immediately eat and then they either [[hoarding (animal behavior)|cache]] or abandon the remainder. The term was invented by Dutch biologist Hans Kruuk after studying [[spotted hyena]]s in [[Africa]]<ref name="Hyena">{{cite book | author= Kruuk, Hans | title= The Spotted Hyena: A study of predation and social behaviour | year= 1972 | isbn= 0-563-20844-9 | page= [https://archive.org/details/velvetclawnatura00macd/page/335 335] | url-access= registration | url= https://archive.org/details/velvetclawnatura00macd/page/335 }}</ref> and [[red fox]]es in [[England]].<ref name="RWTF">{{cite book | author=Macdonald, David | title=Running with the Fox | year=1987 | isbn=0-04-440199-X | page= 224 }}</ref><ref>{{Cite journal|url=https://zslpublications.onlinelibrary.wiley.com/doi/full/10.1111/j.1469-7998.1972.tb04087.x|doi = 10.1111/j.1469-7998.1972.tb04087.x|title = Surplus killing by carnivores|year = 2009|last1 = Kruuk|first1 = Hans|journal = Journal of Zoology|volume = 166|issue = 2|pages = 233–244}}</ref> Some of the other animals which have been observed engaging in surplus killing include [[orcas]],<ref>{{Cite web|date=2015-12-11|title=Suspected surplus killing of harbor seal pups (Phoca vitulina) by killer whales (Orcinus orca)|url=https://www.cascadiaresearch.org/publications/suspected-surplus-killing-harbor-seal-pups-phoca-vitulina-killer-whales-orcinus-orca|access-date=2021-08-09|website=Cascadia Research|language=en}}</ref> [[zooplankton]],<ref>{{Cite journal|last1=Montagnes|first1=David|last2=Fenton|first2=Andy|date=24 September 2012|title=Prey-abundance affects zooplankton assimilation efficiency and the outcome of biogeochemical models|url=https://www.sciencedirect.com/science/article/pii/S0304380012002268|journal=Ecological Modelling|volume=243|pages=1–7|doi=10.1016/j.ecolmodel.2012.05.006}}</ref> [[human]]s,<ref>{{Cite book |title=Living with Coyotes |date=2005-12-31 |url=https://www.degruyter.com/document/doi/10.7560/706323-005/html |work= |pages=42–52 |chapter=Four Surplus Killing |publisher=University of Texas Press |doi=10.7560/706323-005 |isbn=978-0-292-79696-6 |access-date=2022-08-10}}</ref> [[damselfly]] [[naiad (insect)|naiad]]s,<ref>{{Cite journal|last1=Lounibos|first1=L. P.|last2=Makhni|first2=S.|last3=Alto|first3=B. W.|last4=Kesavaraju|first4=B.|date=March 2008|title=Surplus Killing by Predatory Larvae of Corethrella appendiculata: Prepupal Timing and Site-Specific Attack on Mosquito Prey|journal=Journal of Insect Behavior|volume=21|issue=2|pages=47–54|doi=10.1007/s10905-007-9103-2|pmid=19081802|pmc=2600435}}</ref> predaceous [[mite]]s {{CN|date=June 2022}}, [[marten]]s,<ref>{{Cite journal|last1=Brzeziński|first1=Marcin|last2=Rodak|first2=Łukasz|last3=Zalewski|first3=Andrzej|date=2014|title="Reversed" intraguild predation: red fox cubs killed by pine marten|journal=Acta Theriologica|volume=59|issue=3|pages=473–477|doi=10.1007/s13364-014-0179-8|pmid=24954928|pmc=4058055}}</ref> [[weasel]]s,<ref>{{Cite journal|last1=Oksanen|first1=Tarja|last2=Oksanen|first2=Lauri|last3=Fretwell|first3=Stephen D.|date=1985|title=Surplus Killing in the Hunting Strategy of Small Predators|url=https://www.jstor.org/stable/2461358|journal=The American Naturalist|volume=126|issue=3|pages=328–346|doi=10.1086/284420|jstor=2461358|s2cid=84799770|issn=0003-0147}}</ref> [[honey badger]]s,<ref name=":0">{{Cite journal|last1=Shen|first1=Zuolin|last2=Wei|first2=Junjie|date=2018-06-01|title=Hopf bifurcation analysis in a diffusive predator-prey system with delay and surplus killing effect|journal=Mathematical Biosciences and Engineering: MBE|volume=15|issue=3|pages=693–715|doi=10.3934/mbe.2018031|pmid=30380326|doi-access=free}}</ref> [[jaguar]] {{CN|date=June 2022}}, [[leopard]]s,<ref name=":0" /> [[lion]]s,<ref name=":1">{{Cite journal|last=Kruuk|first=Hans|date=1972|title=Surplus killing by carnivores|url=https://zslpublications.onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-7998.1972.tb04087.x|journal=Journal of Zoology|language=en|volume=166|issue=2|pages=233–244|doi=10.1111/j.1469-7998.1972.tb04087.x}}</ref><ref name=":0" /> [[Wolf|wolves]],<ref>{{Cite journal|last1=Miller|first1=Frank L.|last2=Gunn|first2=Anne|last3=Broughton|first3=Eric|date=2011-02-14|title=Surplus killing as exemplified by wolf predation on newborn caribou|url=https://cdnsciencepub.com/doi/abs/10.1139/z85-045|journal=Canadian Journal of Zoology|volume=63|issue=2|pages=295–300|language=en|doi=10.1139/z85-045}}</ref> [[spider]]s,<ref name=":0" /> [[brown bear]]s,<ref>{{Cite journal|last1=Lincoln|first1=Alexandra E|last2=Quinn|first2=Thomas P|date=2019-03-04|title=Optimal foraging or surplus killing: selective consumption and discarding of salmon by brown bears|journal=Behavioral Ecology|volume=30|issue=1|pages=202–212|doi=10.1093/beheco/ary139|doi-access=free}}</ref><ref>{{cite web|url=http://www.gi.alaska.edu/ScienceForum/ASF15/1547.html|archive-url=https://web.archive.org/web/20110807080708/http://www.gi.alaska.edu/ScienceForum/ASF15/1547.html|url-status=dead|archive-date=August 7, 2011|title=Far North Grizzlies Develop Taste for Muskoxen, Alaska Science Forum|date=June 21, 2011}}</ref> [[American black bear]]s,<ref>{{Cite journal|last1=Horstman|first1=Louise P.|last2=Gunson|first2=John R.|date=1982|title=Black Bear Predation on Livestock in Alberta|url=https://www.jstor.org/stable/3781799|journal=Wildlife Society Bulletin |volume=10|issue=1|pages=34–39|jstor=3781799}}</ref> [[polar bear]]s,<ref name=":1" /> [[coyote]]s,<ref name=":1" /><ref>{{Cite book|last=Ewer|first=R. F.|title=Ethology ofmammals|publisher=Logos Press Ltd.|year=1968|location=London}}</ref> [[lynx]]es,<ref>{{Cite journal|last1=Odden|first1=John|last2=Linnell|first2=John D. C.|last3=Moa|first3=Pål Fossland|last4=Herfindal|first4=Ivar|last5=Kvam|first5=Tor|last6=Andersen|first6=Reidar|date=2002|title=Lynx Depredation on Domestic Sheep in Norway|url=https://www.jstor.org/stable/3802876|journal=The Journal of Wildlife Management|volume=66|issue=1|pages=98–105|doi=10.2307/3802876|jstor=3802876}}</ref> [[mink]]s,<ref>{{Cite journal|date=2002-03-01|title=Surplus killing by introduced predators in Australia—evidence for ineffective anti-predator adaptations in native prey species?|url=https://www.sciencedirect.com/science/article/abs/pii/S0006320701001392|journal=Biological Conservation|language=en|volume=103|issue=3|pages=283–301|doi=10.1016/S0006-3207(01)00139-2|last1=Short|first1=Jeff|last2=Kinnear|first2=J.E.|last3=Robley|first3=Alan}}</ref> [[raccoon]]s<ref>{{Cite journal|last1=Hartman|first1=Lisa H.|last2=Gaston|first2=Anthony J.|last3=Eastman|first3=Donald S.|date=1997|title=Raccoon Predation on Ancient Murrelets on East Limestone Island, British Columbia|url=https://www.jstor.org/stable/3802594|journal=The Journal of Wildlife Management|volume=61|issue=2|pages=377–388|doi=10.2307/3802594|jstor=3802594}}</ref> and [[dog]]s.{{CN|date=June 2022}}


'''Surplus killing''', also known as '''excessive killing''', '''henhouse syndrome''',<ref name="LSM148" /><ref>{{cite book|last=Moskowitz|first=David|title=Wolves in the Land of Salmon.|date=4 February 2013|publisher=Timber Press|isbn=978-1604692273|pages=144}}</ref> or '''overkill''',<ref>{{Cite journal|last=Mysterud|first=Ivar|date=1980|title=Bear Management and Sheep Husbandry in Norway, with a Discussion of Predatory Behavior Significant for Evaluation of Livestock Losses|url=http://www.jstor.org/stable/3872873|journal=Bears: Their Biology and Management|volume=4|pages=233–241|doi=10.2307/3872873|jstor=3872873|issn=1936-0614}}</ref> is a common behavior exhibited by [[predator]]s, in which they kill more [[prey]] than they can immediately eat and then they either [[hoarding (animal behavior)|cache]] or abandon the remainder. The term was invented by Dutch biologist Hans Kruuk after studying [[spotted hyena]]s in [[Africa]]<ref name="Hyena">{{cite book | author= Kruuk, Hans | title= The Spotted Hyena: A study of predation and social behaviour | year= 1972 | isbn= 0-563-20844-9 | page= [https://archive.org/details/velvetclawnatura00macd/page/335 335] | publisher= BBC Books | url-access= registration | url= https://archive.org/details/velvetclawnatura00macd/page/335 }}</ref> and [[red fox]]es in [[England]].<ref name="RWTF">{{cite book | author=Macdonald, David | title=Running with the Fox | year=1987 | isbn=0-04-440199-X | page= 224 | publisher=Unwin Hyman }}</ref><ref name="Surplus killing by carnivores">{{Cite journal|url=https://zslpublications.onlinelibrary.wiley.com/doi/full/10.1111/j.1469-7998.1972.tb04087.x|doi = 10.1111/j.1469-7998.1972.tb04087.x|title = Surplus killing by carnivores|year = 2009|last1 = Kruuk|first1 = Hans|journal = Journal of Zoology|volume = 166|issue = 2|pages = 233–244}}</ref>

== Species ==
Some of animals which have been observed engaging in surplus killing hunt alone, some are mainly [[pack hunter]]s. Some opinions include [[human]]s.<ref>{{Cite book |title=Living with Coyotes |date=2005-12-31 |url=https://www.degruyter.com/document/doi/10.7560/706323-005/html |pages=42–52 |chapter=Four Surplus Killing |publisher=University of Texas Press |doi=10.7560/706323-005 |doi-broken-date=2024-04-24 |isbn=978-0-292-79696-6 |s2cid=243782280 |access-date=2022-08-10}}</ref>

Bears: [[brown bear]]s,<ref>{{Cite journal|last1=Lincoln|first1=Alexandra E|last2=Quinn|first2=Thomas P|date=2019-03-04|title=Optimal foraging or surplus killing: selective consumption and discarding of salmon by brown bears|journal=Behavioral Ecology|volume=30|issue=1|pages=202–212|doi=10.1093/beheco/ary139|doi-access=free}}</ref><ref>{{cite web|url=http://www.gi.alaska.edu/ScienceForum/ASF15/1547.html|archive-url=https://web.archive.org/web/20110807080708/http://www.gi.alaska.edu/ScienceForum/ASF15/1547.html|url-status=dead|archive-date=August 7, 2011|title=Far North Grizzlies Develop Taste for Muskoxen, Alaska Science Forum|date=June 21, 2011}}</ref> [[American black bear]]s,<ref>{{Cite journal|last1=Horstman|first1=Louise P.|last2=Gunson|first2=John R.|date=1982|title=Black Bear Predation on Livestock in Alberta|url=https://www.jstor.org/stable/3781799|journal=Wildlife Society Bulletin |volume=10|issue=1|pages=34–39|jstor=3781799}}</ref> [[polar bear]]s,<ref name=":1" />

Large cats: [[jaguar]] {{Citation needed|date=June 2022}}, [[leopard]]s,<ref name=":0" /> [[lion]]s,<ref name=":1">{{Cite journal|last=Kruuk|first=Hans|date=1972|title=Surplus killing by carnivores|url=https://zslpublications.onlinelibrary.wiley.com/doi/abs/10.1111/j.1469-7998.1972.tb04087.x|journal=Journal of Zoology|language=en|volume=166|issue=2|pages=233–244|doi=10.1111/j.1469-7998.1972.tb04087.x}}</ref><ref name=":0" /> [[lynx]]es,<ref>{{Cite journal|last1=Odden|first1=John|last2=Linnell|first2=John D. C.|last3=Moa|first3=Pål Fossland|last4=Herfindal|first4=Ivar|last5=Kvam|first5=Tor|last6=Andersen|first6=Reidar|date=2002|title=Lynx Depredation on Domestic Sheep in Norway|url=https://www.jstor.org/stable/3802876|journal=The Journal of Wildlife Management|volume=66|issue=1|pages=98–105|doi=10.2307/3802876|jstor=3802876}}</ref>

Canines: [[Wolf|wolves]],<ref>{{Cite journal|last1=Miller|first1=Frank L.|last2=Gunn|first2=Anne|last3=Broughton|first3=Eric|date=2011-02-14|title=Surplus killing as exemplified by wolf predation on newborn caribou|url=https://cdnsciencepub.com/doi/abs/10.1139/z85-045|journal=Canadian Journal of Zoology|volume=63|issue=2|pages=295–300|language=en|doi=10.1139/z85-045}}</ref> [[coyote]]s,<ref name=":1" /><ref>{{Cite book|last=Ewer|first=R. F.|title=Ethology ofmammals|publisher=Logos Press Ltd.|year=1968|location=London}}</ref> [[red fox]],<ref name="RWTF"/><ref name="Surplus killing by carnivores"/> [[dog]]s.{{Citation needed|date=June 2022}}

Others: [[marten]]s,<ref>{{Cite journal|last1=Brzeziński|first1=Marcin|last2=Rodak|first2=Łukasz|last3=Zalewski|first3=Andrzej|date=2014|title="Reversed" intraguild predation: red fox cubs killed by pine marten|journal=Acta Theriologica|volume=59|issue=3|pages=473–477|doi=10.1007/s13364-014-0179-8|pmid=24954928|pmc=4058055}}</ref> [[weasel]]s,<ref>{{Cite journal|last1=Oksanen|first1=Tarja|last2=Oksanen|first2=Lauri|last3=Fretwell|first3=Stephen D.|date=1985|title=Surplus Killing in the Hunting Strategy of Small Predators|url=https://www.jstor.org/stable/2461358|journal=The American Naturalist|volume=126|issue=3|pages=328–346|doi=10.1086/284420|jstor=2461358|s2cid=84799770|issn=0003-0147}}</ref> [[honey badger]]s,<ref name=":0">{{Cite journal|last1=Shen|first1=Zuolin|last2=Wei|first2=Junjie|date=2018-06-01|title=Hopf bifurcation analysis in a diffusive predator-prey system with delay and surplus killing effect|journal=Mathematical Biosciences and Engineering: MBE|volume=15|issue=3|pages=693–715|doi=10.3934/mbe.2018031|pmid=30380326|doi-access=free}}</ref> [[mink]]s,<ref>{{Cite journal|date=2002-03-01|title=Surplus killing by introduced predators in Australia—evidence for ineffective anti-predator adaptations in native prey species?|url=https://www.sciencedirect.com/science/article/abs/pii/S0006320701001392|journal=Biological Conservation|language=en|volume=103|issue=3|pages=283–301|doi=10.1016/S0006-3207(01)00139-2|last1=Short|first1=Jeff|last2=Kinnear|first2=J.E.|last3=Robley|first3=Alan|bibcode=2002BCons.103..283S }}</ref> [[raccoon]]s,<ref>{{Cite journal|last1=Hartman|first1=Lisa H.|last2=Gaston|first2=Anthony J.|last3=Eastman|first3=Donald S.|date=1997|title=Raccoon Predation on Ancient Murrelets on East Limestone Island, British Columbia|url=https://www.jstor.org/stable/3802594|journal=The Journal of Wildlife Management|volume=61|issue=2|pages=377–388|doi=10.2307/3802594|jstor=3802594}}</ref> [[spotted hyena]],<ref name="Hyena"/> [[orcas]]<ref>{{Cite web|date=2015-12-11|title=Suspected surplus killing of harbor seal pups (Phoca vitulina) by killer whales (Orcinus orca)|url=https://www.cascadiaresearch.org/publications/suspected-surplus-killing-harbor-seal-pups-phoca-vitulina-killer-whales-orcinus-orca|access-date=2021-08-09|website=Cascadia Research|language=en}}</ref>

Invertebrae: [[zooplankton]],<ref>{{Cite journal|last1=Montagnes|first1=David|last2=Fenton|first2=Andy|date=24 September 2012|title=Prey-abundance affects zooplankton assimilation efficiency and the outcome of biogeochemical models|url=https://www.sciencedirect.com/science/article/pii/S0304380012002268|journal=Ecological Modelling|volume=243|pages=1–7|doi=10.1016/j.ecolmodel.2012.05.006}}</ref> [[damselfly]] [[naiad (insect)|naiad]]s,<ref>{{Cite journal|last1=Lounibos|first1=L. P.|last2=Makhni|first2=S.|last3=Alto|first3=B. W.|last4=Kesavaraju|first4=B.|date=March 2008|title=Surplus Killing by Predatory Larvae of Corethrella appendiculata: Prepupal Timing and Site-Specific Attack on Mosquito Prey|journal=Journal of Insect Behavior|volume=21|issue=2|pages=47–54|doi=10.1007/s10905-007-9103-2|pmid=19081802|pmc=2600435|bibcode=2008JIBeh..21...47L }}</ref> predaceous [[mite]]s {{Citation needed|date=June 2022}}, [[spider]]s,<ref name=":0" />

== Example cases ==
In [[Tasmania]], in a single dog attack, 58 [[little penguin]]s were killed.<ref>{{Cite web|url=https://www.upi.com/Top_News/World-News/2018/10/17/Dog-attack-kill-58-penguins-in-Tasmania/9631539800130/|title=Dog attack kills 58 penguins in Tasmania|last=Adamczyk|first=Ed|date=2018-10-17|website=UPI|language=en-AU|access-date=2020-04-04}}</ref> In [[mainland Australia]], a single fox once killed around 74 penguins over several days, eating almost nothing.<ref>{{Cite web|url=https://www.abc.net.au/news/2017-08-10/penguins-killed-by-fox-victoria-middle-island/8794828|title=Penguins killed in fox attack on Victoria's Middle Island|last=Peacock|first=Sue|date=2017-08-10|website=ABC News|language=en-AU|access-date=2019-09-26}}</ref> One leopard in [[Cape Province]], [[South Africa]] killed 51 sheep and lambs in a single incident.<ref>{{cite journal |last1=Stuart |first1=C. T. |year=1986 |title=The incidence of surplus killing by Panthera pardus and Felis caracal in Cape Province, South Africa |journal=Mammalia |volume=50 |issue=4 |pages=556–558 |issn=0025-1461 | doi = 10.1515/mamm.1986.50.4.553 }}</ref> Similarly, two [[caracal]] in Cape Province killed 22 sheep in one night, eating only part of the buttock of one carcass.<ref>{{cite journal |last1=Skinner |first1=J. D. |year=1979 |title=Feeding behaviour in Caracal ''Felis caracal'' |journal=Journal of Zoology |volume=189 |issue=4 |pages=523–525 | doi = 10.1111/j.1469-7998.1979.tb03979.x }}</ref> Up to 19 spotted hyenas once killed 82 [[Thomson's gazelle]] and badly injured 27, eating just 16%.<ref name=LSM148>{{cite book|last=Mills|first=L. Scott|title=Conservation of wildlife populations: demography, genetics, and management|date=17 December 2012|publisher=Wiley-Blackwell|location=Hoboken, NJ|isbn=9780470671504|pages=148|edition=2nd}}</ref>
In [[Tasmania]], in a single dog attack, 58 [[little penguin]]s were killed.<ref>{{Cite web|url=https://www.upi.com/Top_News/World-News/2018/10/17/Dog-attack-kill-58-penguins-in-Tasmania/9631539800130/|title=Dog attack kills 58 penguins in Tasmania|last=Adamczyk|first=Ed|date=2018-10-17|website=UPI|language=en-AU|access-date=2020-04-04}}</ref> In [[mainland Australia]], a single fox once killed around 74 penguins over several days, eating almost nothing.<ref>{{Cite web|url=https://www.abc.net.au/news/2017-08-10/penguins-killed-by-fox-victoria-middle-island/8794828|title=Penguins killed in fox attack on Victoria's Middle Island|last=Peacock|first=Sue|date=2017-08-10|website=ABC News|language=en-AU|access-date=2019-09-26}}</ref> One leopard in [[Cape Province]], [[South Africa]] killed 51 sheep and lambs in a single incident.<ref>{{cite journal |last1=Stuart |first1=C. T. |year=1986 |title=The incidence of surplus killing by Panthera pardus and Felis caracal in Cape Province, South Africa |journal=Mammalia |volume=50 |issue=4 |pages=556–558 |issn=0025-1461 | doi = 10.1515/mamm.1986.50.4.553 }}</ref> Similarly, two [[caracal]] in Cape Province killed 22 sheep in one night, eating only part of the buttock of one carcass.<ref>{{cite journal |last1=Skinner |first1=J. D. |year=1979 |title=Feeding behaviour in Caracal ''Felis caracal'' |journal=Journal of Zoology |volume=189 |issue=4 |pages=523–525 | doi = 10.1111/j.1469-7998.1979.tb03979.x }}</ref> Up to 19 spotted hyenas once killed 82 [[Thomson's gazelle]] and badly injured 27, eating just 16%.<ref name=LSM148>{{cite book|last=Mills|first=L. Scott|title=Conservation of wildlife populations: demography, genetics, and management|date=17 December 2012|publisher=Wiley-Blackwell|location=Hoboken, NJ|isbn=9780470671504|pages=148|edition=2nd}}</ref>


In late autumn, [[least weasel]]s often surplus-kill [[vole]] and then dig them up and eat them on winter days when it is too cold to hunt. In March 2016, a pack of 9 grey wolves in Wyoming was found to have killed 19 [[elk]]. John Lund, of the Wyoming Game and Fish Department, said that he had never documented surplus killings that extreme from grey wolves.<ref>{{cite web|url=http://www.calgarysun.com/2016/03/25/wyoming-wolf-pack-kills-19-elk-in-rare-surplus-killing|title=Wyoming wolf pack kills 19 elk in rare 'surplus killing'|access-date=2016-03-25|archive-url=https://web.archive.org/web/20160404194628/http://www.calgarysun.com/2016/03/25/wyoming-wolf-pack-kills-19-elk-in-rare-surplus-killing|archive-date=2016-04-04|url-status=dead}}</ref> In Florida, laboratory experiments documented cases of surplus killing in larvae of the predatory midge ''[[Corethrella]] appendiculata'' against specific larval stages of different species of mosquitoes of the genus ''[[Toxorhynchites]]''.<ref>{{cite journal|author=L.P. Lounibos, S. Mahkni, B.W.Alto, B. Kesavaraju|title=Surplus Killing by Predatory Larvae of ''Corethrella appendiculata'': Prepupal Timing and Site-Specific Attack on Mosquito Prey | journal=Journal of Insects Behaviour|volume=21 |number=2 |pages=47–54|date=Mar 2008 |doi= 10.1007/s10905-007-9103-2 |pmid=19081802 |pmc=2600435 }}</ref>
In late autumn, [[least weasel]]s often surplus-kill [[vole]] and then dig them up and eat them on winter days when it is too cold to hunt. In March 2016, a pack of 9 grey wolves in Wyoming was found to have killed 19 [[elk]]. John Lund, of the Wyoming Game and Fish Department, said that he had never documented surplus killings that extreme from grey wolves.<ref>{{cite web|url=http://www.calgarysun.com/2016/03/25/wyoming-wolf-pack-kills-19-elk-in-rare-surplus-killing|title=Wyoming wolf pack kills 19 elk in rare 'surplus killing'|access-date=2016-03-25|archive-url=https://web.archive.org/web/20160404194628/http://www.calgarysun.com/2016/03/25/wyoming-wolf-pack-kills-19-elk-in-rare-surplus-killing|archive-date=2016-04-04|url-status=dead}}</ref> In Florida, laboratory experiments documented cases of surplus killing in larvae of the predatory midge ''[[Corethrella]] appendiculata'' against specific larval stages of different species of mosquitoes of the genus ''[[Toxorhynchites]]''.<ref>{{cite journal|author=L.P. Lounibos, S. Mahkni, B.W.Alto, B. Kesavaraju|title=Surplus Killing by Predatory Larvae of ''Corethrella appendiculata'': Prepupal Timing and Site-Specific Attack on Mosquito Prey | journal=Journal of Insect Behavior|volume=21 |number=2 |pages=47–54|date=Mar 2008 |doi= 10.1007/s10905-007-9103-2 |pmid=19081802 |pmc=2600435 |bibcode=2008JIBeh..21...47L }}</ref>


== Possible causes ==
In surplus killing, predators eat only the most-preferred animals and animal parts. Bears engaging in surplus killing of [[salmon]] are more likely to eat unspawned fish because of higher muscle quality, and high-energy parts such as brains and eggs.<ref name=LSM148/> Surplus killing can deplete the overall [[food supply]], waste predator energy and risk them being injured. Nonetheless, researchers say animals surplus-kill whenever they can, in order to procure food for offspring and others, to gain valuable killing experience, and to create the opportunity to eat the carcass later when they are hungry again.<ref name=LSM148/><ref>{{cite book|last=Hansen|first=Kevin|title=Bobcat: master of survival|url=https://archive.org/details/bobcatmastersurv00hans|url-access=limited|year=2006|publisher=Oxford University Press|location=New York|isbn=0195183037|pages=[https://archive.org/details/bobcatmastersurv00hans/page/n130 114]|edition=[Online-Ausg.]}}</ref>
In surplus killing, predators eat only the most-preferred animals and animal parts. Bears engaging in surplus killing of [[salmon]] are more likely to eat unspawned fish because of higher muscle quality, and high-energy parts such as brains and eggs.<ref name=LSM148/> Surplus killing can deplete the overall [[food supply]], waste predator energy and risk their being injured. Nonetheless, researchers say animals surplus-kill whenever they can, in order to procure food for offspring and others, to gain valuable killing experience, and to create the opportunity to eat the carcass later when they are hungry again.<ref name=LSM148/><ref>{{cite book|last=Hansen|first=Kevin|title=Bobcat: master of survival|url=https://archive.org/details/bobcatmastersurv00hans|url-access=limited|year=2006|publisher=Oxford University Press|location=New York|isbn=0195183037|pages=[https://archive.org/details/bobcatmastersurv00hans/page/n130 114]|edition=[Online-Ausg.]}}</ref>


== See also ==
== See also ==
Line 18: Line 34:
==Bibliography==
==Bibliography==
* Jennifer L. Maupin and Susan Reichert, ''[http://beheco.oxfordjournals.org/cgi/content/full/12/5/569 Superfluous killing in spiders]''.
* Jennifer L. Maupin and Susan Reichert, ''[http://beheco.oxfordjournals.org/cgi/content/full/12/5/569 Superfluous killing in spiders]''.
* Joseph K. Gaydos, Stephen Raverty, Robin W. Baird, and Richard W. Osborne, ''[http://www.cascadiaresearch.org/robin/GaydosetalNWN2005.pdf#search=%22surplus%20killing%22 SUSPECTED SURPLUS KILLING OF HARBOR SEAL PUPS (PHOCA VITULINA) BY KILLER WHALES (ORCINUS ORCA)]''.
* Joseph K. Gaydos, Stephen Raverty, Robin W. Baird, and Richard W. Osborne, ''[http://www.cascadiaresearch.org/robin/GaydosetalNWN2005.pdf#search=%22surplus%20killing%22 SUSPECTED SURPLUS KILLING OF HARBOR SEAL PUPS (PHOCA VITULINA) BY KILLER WHALES (ORCINUS ORCA)] {{Webarchive|url=https://web.archive.org/web/20110929115317/http://www.cascadiaresearch.org/robin/GaydosetalNWN2005.pdf#search=%22surplus%20killing%22 |date=2011-09-29 }}''.
* William G. George and Timothy Kimmel, ''[https://web.archive.org/web/20110617101214/http://elibrary.unm.edu/sora/Auk/v094n04/p0782-p0783.pdf#search=%22surplus%20killing%22 A Slaughter of Mice by Common Crows]''.
* William G. George and Timothy Kimmel, ''[https://web.archive.org/web/20110617101214/http://elibrary.unm.edu/sora/Auk/v094n04/p0782-p0783.pdf#search=%22surplus%20killing%22 A Slaughter of Mice by Common Crows]''.
* Wildlife Online: Foxes-Surplus Killing, ''[http://www.wildlifeonline.me.uk/questions_answers_foxes.html#kill Why do foxes kill to excess...]''.
* Wildlife Online: Foxes-Surplus Killing, ''[http://www.wildlifeonline.me.uk/questions_answers_foxes.html#kill Why do foxes kill to excess...]''.
* For Wolves: Ralph Maughan Wolf Report, ''[http://www.forwolves.org/ralph/jacksonwolves7-.htm Jackson Trio makes some surplus kills]''.
* For Wolves: Ralph Maughan Wolf Report, ''[http://www.forwolves.org/ralph/jacksonwolves7-.htm Jackson Trio makes some surplus kills] {{Webarchive|url=https://web.archive.org/web/20060907172227/http://www.forwolves.org/ralph/jacksonwolves7-.htm |date=2006-09-07 }}''.
* High Country News, Zachary Smith, ''[http://www.hcn.org/servlets/hcn.Article?article_id=15048 Wolf pack wiped out for ‘surplus killing’]''.
* High Country News, Zachary Smith, ''[http://www.hcn.org/servlets/hcn.Article?article_id=15048 Wolf pack wiped out for ‘surplus killing’] {{Webarchive|url=https://web.archive.org/web/20060822081003/http://www.hcn.org/servlets/hcn.Article?article_id=15048 |date=2006-08-22 }}''.
* Victor Van Ballenberghe, ''[https://web.archive.org/web/20060917142923/http://wolfsongalaska.org/tech_info.html Technical Information on Wolf Ecology and Wolf/Prey Relationships]''.
* Victor Van Ballenberghe, ''[https://web.archive.org/web/20060917142923/http://wolfsongalaska.org/tech_info.html Technical Information on Wolf Ecology and Wolf/Prey Relationships]''.
* Pierre-Yves Daoust, Andrew Boyne, Ted D’Eon, ''[https://web.archive.org/web/20070708185226/http://wildlife1.usask.ca/newsletters/newsletter9-2.htm#terns Surplus killing of Roseate Terns and Common Terns by a mink]''.
* Pierre-Yves Daoust, Andrew Boyne, Ted D’Eon, ''[https://web.archive.org/web/20070708185226/http://wildlife1.usask.ca/newsletters/newsletter9-2.htm#terns Surplus killing of Roseate Terns and Common Terns by a mink]''.

Latest revision as of 23:10, 17 June 2024

A stoat surplus killing chipmunks (Ernest Thompson Seton, 1909)
Multiple sheep killed by a cougar

Surplus killing, also known as excessive killing, henhouse syndrome,[1][2] or overkill,[3] is a common behavior exhibited by predators, in which they kill more prey than they can immediately eat and then they either cache or abandon the remainder. The term was invented by Dutch biologist Hans Kruuk after studying spotted hyenas in Africa[4] and red foxes in England.[5][6]

Species[edit]

Some of animals which have been observed engaging in surplus killing hunt alone, some are mainly pack hunters. Some opinions include humans.[7]

Bears: brown bears,[8][9] American black bears,[10] polar bears,[11]

Large cats: jaguar [citation needed], leopards,[12] lions,[11][12] lynxes,[13]

Canines: wolves,[14] coyotes,[11][15] red fox,[5][6] dogs.[citation needed]

Others: martens,[16] weasels,[17] honey badgers,[12] minks,[18] raccoons,[19] spotted hyena,[4] orcas[20]

Invertebrae: zooplankton,[21] damselfly naiads,[22] predaceous mites [citation needed], spiders,[12]

Example cases[edit]

In Tasmania, in a single dog attack, 58 little penguins were killed.[23] In mainland Australia, a single fox once killed around 74 penguins over several days, eating almost nothing.[24] One leopard in Cape Province, South Africa killed 51 sheep and lambs in a single incident.[25] Similarly, two caracal in Cape Province killed 22 sheep in one night, eating only part of the buttock of one carcass.[26] Up to 19 spotted hyenas once killed 82 Thomson's gazelle and badly injured 27, eating just 16%.[1]

In late autumn, least weasels often surplus-kill vole and then dig them up and eat them on winter days when it is too cold to hunt. In March 2016, a pack of 9 grey wolves in Wyoming was found to have killed 19 elk. John Lund, of the Wyoming Game and Fish Department, said that he had never documented surplus killings that extreme from grey wolves.[27] In Florida, laboratory experiments documented cases of surplus killing in larvae of the predatory midge Corethrella appendiculata against specific larval stages of different species of mosquitoes of the genus Toxorhynchites.[28]

Possible causes[edit]

In surplus killing, predators eat only the most-preferred animals and animal parts. Bears engaging in surplus killing of salmon are more likely to eat unspawned fish because of higher muscle quality, and high-energy parts such as brains and eggs.[1] Surplus killing can deplete the overall food supply, waste predator energy and risk their being injured. Nonetheless, researchers say animals surplus-kill whenever they can, in order to procure food for offspring and others, to gain valuable killing experience, and to create the opportunity to eat the carcass later when they are hungry again.[1][29]

See also[edit]

References[edit]

  1. ^ a b c d Mills, L. Scott (17 December 2012). Conservation of wildlife populations: demography, genetics, and management (2nd ed.). Hoboken, NJ: Wiley-Blackwell. p. 148. ISBN 9780470671504.
  2. ^ Moskowitz, David (4 February 2013). Wolves in the Land of Salmon. Timber Press. p. 144. ISBN 978-1604692273.
  3. ^ Mysterud, Ivar (1980). "Bear Management and Sheep Husbandry in Norway, with a Discussion of Predatory Behavior Significant for Evaluation of Livestock Losses". Bears: Their Biology and Management. 4: 233–241. doi:10.2307/3872873. ISSN 1936-0614. JSTOR 3872873.
  4. ^ a b Kruuk, Hans (1972). The Spotted Hyena: A study of predation and social behaviour. BBC Books. p. 335. ISBN 0-563-20844-9.
  5. ^ a b Macdonald, David (1987). Running with the Fox. Unwin Hyman. p. 224. ISBN 0-04-440199-X.
  6. ^ a b Kruuk, Hans (2009). "Surplus killing by carnivores". Journal of Zoology. 166 (2): 233–244. doi:10.1111/j.1469-7998.1972.tb04087.x.
  7. ^ "Four Surplus Killing". Living with Coyotes. University of Texas Press. 2005-12-31. pp. 42–52. doi:10.7560/706323-005 (inactive 2024-04-24). ISBN 978-0-292-79696-6. S2CID 243782280. Retrieved 2022-08-10.{{cite book}}: CS1 maint: DOI inactive as of April 2024 (link)
  8. ^ Lincoln, Alexandra E; Quinn, Thomas P (2019-03-04). "Optimal foraging or surplus killing: selective consumption and discarding of salmon by brown bears". Behavioral Ecology. 30 (1): 202–212. doi:10.1093/beheco/ary139.
  9. ^ "Far North Grizzlies Develop Taste for Muskoxen, Alaska Science Forum". June 21, 2011. Archived from the original on August 7, 2011.
  10. ^ Horstman, Louise P.; Gunson, John R. (1982). "Black Bear Predation on Livestock in Alberta". Wildlife Society Bulletin. 10 (1): 34–39. JSTOR 3781799.
  11. ^ a b c Kruuk, Hans (1972). "Surplus killing by carnivores". Journal of Zoology. 166 (2): 233–244. doi:10.1111/j.1469-7998.1972.tb04087.x.
  12. ^ a b c d Shen, Zuolin; Wei, Junjie (2018-06-01). "Hopf bifurcation analysis in a diffusive predator-prey system with delay and surplus killing effect". Mathematical Biosciences and Engineering: MBE. 15 (3): 693–715. doi:10.3934/mbe.2018031. PMID 30380326.
  13. ^ Odden, John; Linnell, John D. C.; Moa, Pål Fossland; Herfindal, Ivar; Kvam, Tor; Andersen, Reidar (2002). "Lynx Depredation on Domestic Sheep in Norway". The Journal of Wildlife Management. 66 (1): 98–105. doi:10.2307/3802876. JSTOR 3802876.
  14. ^ Miller, Frank L.; Gunn, Anne; Broughton, Eric (2011-02-14). "Surplus killing as exemplified by wolf predation on newborn caribou". Canadian Journal of Zoology. 63 (2): 295–300. doi:10.1139/z85-045.
  15. ^ Ewer, R. F. (1968). Ethology ofmammals. London: Logos Press Ltd.
  16. ^ Brzeziński, Marcin; Rodak, Łukasz; Zalewski, Andrzej (2014). ""Reversed" intraguild predation: red fox cubs killed by pine marten". Acta Theriologica. 59 (3): 473–477. doi:10.1007/s13364-014-0179-8. PMC 4058055. PMID 24954928.
  17. ^ Oksanen, Tarja; Oksanen, Lauri; Fretwell, Stephen D. (1985). "Surplus Killing in the Hunting Strategy of Small Predators". The American Naturalist. 126 (3): 328–346. doi:10.1086/284420. ISSN 0003-0147. JSTOR 2461358. S2CID 84799770.
  18. ^ Short, Jeff; Kinnear, J.E.; Robley, Alan (2002-03-01). "Surplus killing by introduced predators in Australia—evidence for ineffective anti-predator adaptations in native prey species?". Biological Conservation. 103 (3): 283–301. Bibcode:2002BCons.103..283S. doi:10.1016/S0006-3207(01)00139-2.
  19. ^ Hartman, Lisa H.; Gaston, Anthony J.; Eastman, Donald S. (1997). "Raccoon Predation on Ancient Murrelets on East Limestone Island, British Columbia". The Journal of Wildlife Management. 61 (2): 377–388. doi:10.2307/3802594. JSTOR 3802594.
  20. ^ "Suspected surplus killing of harbor seal pups (Phoca vitulina) by killer whales (Orcinus orca)". Cascadia Research. 2015-12-11. Retrieved 2021-08-09.
  21. ^ Montagnes, David; Fenton, Andy (24 September 2012). "Prey-abundance affects zooplankton assimilation efficiency and the outcome of biogeochemical models". Ecological Modelling. 243: 1–7. doi:10.1016/j.ecolmodel.2012.05.006.
  22. ^ Lounibos, L. P.; Makhni, S.; Alto, B. W.; Kesavaraju, B. (March 2008). "Surplus Killing by Predatory Larvae of Corethrella appendiculata: Prepupal Timing and Site-Specific Attack on Mosquito Prey". Journal of Insect Behavior. 21 (2): 47–54. Bibcode:2008JIBeh..21...47L. doi:10.1007/s10905-007-9103-2. PMC 2600435. PMID 19081802.
  23. ^ Adamczyk, Ed (2018-10-17). "Dog attack kills 58 penguins in Tasmania". UPI. Retrieved 2020-04-04.
  24. ^ Peacock, Sue (2017-08-10). "Penguins killed in fox attack on Victoria's Middle Island". ABC News. Retrieved 2019-09-26.
  25. ^ Stuart, C. T. (1986). "The incidence of surplus killing by Panthera pardus and Felis caracal in Cape Province, South Africa". Mammalia. 50 (4): 556–558. doi:10.1515/mamm.1986.50.4.553. ISSN 0025-1461.
  26. ^ Skinner, J. D. (1979). "Feeding behaviour in Caracal Felis caracal". Journal of Zoology. 189 (4): 523–525. doi:10.1111/j.1469-7998.1979.tb03979.x.
  27. ^ "Wyoming wolf pack kills 19 elk in rare 'surplus killing'". Archived from the original on 2016-04-04. Retrieved 2016-03-25.
  28. ^ L.P. Lounibos, S. Mahkni, B.W.Alto, B. Kesavaraju (Mar 2008). "Surplus Killing by Predatory Larvae of Corethrella appendiculata: Prepupal Timing and Site-Specific Attack on Mosquito Prey". Journal of Insect Behavior. 21 (2): 47–54. Bibcode:2008JIBeh..21...47L. doi:10.1007/s10905-007-9103-2. PMC 2600435. PMID 19081802.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  29. ^ Hansen, Kevin (2006). Bobcat: master of survival ([Online-Ausg.] ed.). New York: Oxford University Press. pp. 114. ISBN 0195183037.

Bibliography[edit]

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