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{{Short description|Predator at the top of a food chain}}
{{Other uses}}
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[[File:020 The lion king Snyggve in the Serengeti National Park Photo by Giles Laurent.jpg|thumb|The [[lion]] is the world's [[List of largest cats|second-largest]] big cat and serves as an apex land predator in Africa.<ref name="OrdizBischofSwenson2013">{{cite journal |last1=Ordiz |first1=Andrés |last2=Bischof |first2=Richard |last3=Swenson |first3=Jon E. |title=Saving large carnivores, but losing the apex predator? |journal=Biological Conservation |volume=168 |year=2013 |doi=10.1016/j.biocon.2013.09.024 |pages=128–133|doi-access=free |bibcode=2013BCons.168..128O |hdl=11250/2492589 |hdl-access=free }}</ref><ref>{{Cite journal |last1=Kristoffer T. Everatt |last2=Jennifer F. Moore |last3=Graham I.H. Kerley |date=2019 |title=Africa's apex predator, the lion, is limited by interference and exploitive competition with humans |url=https://search.nal.usda.gov/discovery/search?query=lds35,contains,6594565-01nal_inst,AND&tab=LibraryCatalog&search_scope=MyInstitution&vid=01NAL_INST:MAIN&mode=advanced&offset=0 |journal=Global Ecology and Conservation |volume=20 |language=English |pages=e00758 |doi=10.1016/j.gecco.2019.e00758 |s2cid=202023472 |issn=2351-9894|doi-access=free |bibcode=2019GEcoC..2000758E }}</ref>]]
[[File:SaltwaterCrocodile('Maximo').jpg|thumb|The [[saltwater crocodile]] is the largest living reptile and the dominant predator throughout its range.<ref>Whiting, Frances. [http://www.australiazoo.com/about-us/in-the-news/index.php?news=393 "Terri fights to halt croc eggs harvest."] {{webarchive|url=https://web.archive.org/web/20101028052754/http://australiazoo.com/about-us/in-the-news/index.php?news=393 |date=2010-10-28 }} ''Australia Zoo''. 2007-06-11. Retrieved 2010-01-25.</ref><ref>[http://animals.nationalgeographic.com/animals/reptiles/saltwater-crocodile.html "Saltwater Crocodile."] {{webarchive|url=https://web.archive.org/web/20130906074746/http://animals.nationalgeographic.com/animals/reptiles/saltwater-crocodile.html |date=2013-09-06 }} ''National Geographic''. Retrieved 2010-01-25.</ref>]]
[[File:Comparison of size of orca and great white shark.svg|thumb|The [[great white shark]] (bottom) is one of the top marine predators; however, the [[orca]] (top) is known to prey upon them. Thus, great white sharks are not true apex predators. Despite coexisting the majority of the time, confrontations may occur in stressed environments when both species compete for limited resources.]]
An '''apex predator''', also known as a '''top predator''' or '''superpredator''', is a [[predator]]{{efn|[[Zoologist]]s generally exclude [[Parasitism|parasites]] from trophic levels as they are (often much) smaller than their hosts, and individual species with multiple hosts at different life-cycle stages would occupy multiple levels. Otherwise they would often be at the top level, above apex predators.<ref name="Sukhdeo2012">{{cite journal |last=Sukhdeo |first=Michael V. K. |title=Where are the parasites in food webs? |journal=Parasites & Vectors |volume=5 |issue=1 |year=2012 |doi=10.1186/1756-3305-5-239 |pmid=23092160 |pmc=3523981 |page=239 |doi-access=free }}</ref>}} at the top of a [[food chain]], without natural predators of its own.<ref>{{cite web |url=http://www.etymonline.com/index.php?search=predation&searchmode=none |title=predator |publisher=Online Etymological Dictionary |access-date=2010-01-25 |url-status=live |archive-url=https://web.archive.org/web/20090701025327/http://www.etymonline.com/index.php?search=predation&searchmode=none |archive-date=2009-07-01 }}</ref><ref>{{cite web |url=https://www.pbs.org/kqed/oceanadventures/glossary/ |title=apex predator |publisher=PBS |access-date=2010-01-25 |url-status=live |archive-url=https://web.archive.org/web/20090722012639/http://www.pbs.org/kqed/oceanadventures/glossary/ |archive-date=2009-07-22}}</ref>

Apex predators are usually defined in terms of [[trophic dynamics]], meaning that they occupy the highest [[trophic level]]s. Food chains are often far shorter on land, usually limited to being secondary consumers – for example, [[Gray wolf|wolves]] prey mostly upon large [[herbivore]]s (primary consumers), which eat plants (primary producers). The apex predator concept is applied in [[wildlife management]], [[conservation biology|conservation]], and [[ecotourism]].

Apex predators have a long evolutionary history, dating at least to the [[Cambrian]] period when animals such as ''[[Anomalocaris]]'' and ''[[Timorebestia]]'' dominated the seas.

[[Human]]s have for many centuries interacted with other apex predators including the [[wolf]], [[Bird of prey|birds of prey]], and [[cormorant]]s to hunt game animals, birds, and fish respectively. More recently, humans have started interacting with apex predators in new ways. These include interactions via [[ecotourism]], such as with the [[tiger shark]], and through [[rewilding (conservation biology)|rewilding]] efforts, such as the proposed reintroduction of the [[Iberian lynx]].

==Ecological roles==

===Effects on community===
[[File:Skua Runde.jpg|thumb|right|The [[great skua]] is an aerial apex predator, both preying on other seabirds and [[Kleptoparasitism|bullying them for their catches]].<ref name="Leat Bourgeon Eze Muir pp. 569–576">{{cite journal |last1=Leat |first1=Eliza H. K. |last2=Bourgeon |first2=Sophie |last3=Eze |first3=Jude I. |last4=Muir |first4=Derek C.G. |last5=Williamson |first5=Mary |last6=Bustnes |first6=Jan O. |last7=Furness |first7=Robert W. |last8=Borgå |first8=Katrine |title=Perfluoroalkyl substances in eggs and plasma of an avian top predator, great skua (Stercorarius skua), in the north Atlantic |journal=Environmental Toxicology and Chemistry |volume=32 |issue=3 |date=2013-02-15 |doi=10.1002/etc.2101 |pmid=23258709 |pages=569–576|doi-access=free }}</ref>]]

Apex [[predator]]s affect prey species' [[population dynamics]] and populations of other predators, both in aquatic and terrestrial ecosystems. Non-native predatory fish, for instance, have sometimes devastated formerly dominant predators. A lake manipulation study found that when the non-native [[smallmouth bass]] was removed, [[lake trout]], the suppressed native apex predator, diversified its prey selection and increased its [[trophic level]].<ref>{{cite journal |author1=Lepak, Jesse M. |author2=Kraft, Clifford E. |author3=Weidel, Brian C. |date=March 2006 |title=Rapid food web recovery in response to removal of an introduced apex predator |url=http://www.dnr.cornell.edu/cek7/Publications/Lepak_et_al_2006.pdf |journal=Canadian Journal of Fisheries and Aquatic Sciences |volume=63 |issue=3 |pages=569–575 |doi=10.1139/f05-248 |archive-url=https://web.archive.org/web/20080911091513/http://www.dnr.cornell.edu/cek7/Publications/Lepak_et_al_2006.pdf |archive-date=11 September 2008}}</ref> As a terrestrial example, the [[badger]], an apex predator, preys upon and also competes with the [[hedgehog]], a [[mesopredator]], for food such as insects, small mammals, reptiles, amphibians, and the eggs of ground-nesting birds. Removal of badgers (in a trial investigating [[bovine tuberculosis]]) caused hedgehog densities to more than double.<ref name="Trewby 2014">{{cite journal |last1=Trewby |first1=Iain D. |last2=Young |first2=Richard |last3=McDonald |first3=Robbie A. |last4=Wilson |first4=Gavin J. |last5=Davison |first5=John |last6=Walker |first6=Neil |last7=Robertson |first7=Andrew |last8=Doncaster |first8=C. Patrick |last9=Delahay |first9=Richard J. |date=April 2014 |editor-last=Criscuolo |editor-first=François |title=Impacts of Removing Badgers on Localised Counts of Hedgehogs |journal=PLOS ONE |volume=9 |issue=4 |page=e95477 |bibcode=2014PLoSO...995477T |doi=10.1371/journal.pone.0095477 |pmc=3988185 |pmid=24736454 |doi-access=free}}</ref> Predators that exert top-down control on organisms in their community are often considered [[keystone species]].<ref name="Davic2003">{{cite journal |author=Davic, Robert D. |year=2003 |title=Linking Keystone Species and Functional Groups: A New Operational Definition of the Keystone Species Concept |url=http://www.consecol.org/vol7/iss1/resp11/ |journal=Conservation Ecology |volume=7 |doi=10.5751/ES-00502-0701r11 |access-date=2011-02-03 |hdl=10535/2966|hdl-access=free }}</ref> 
<!--HUMANS are discussed in a separate section below-->

===Effects on ecosystem===
{{Further |Trophic cascade}}

Apex predators can have profound effects on ecosystems, as the consequences of both controlling prey density and restricting smaller predators, and may be capable of self-regulation.<ref name="WallachIzhaki2015">{{cite journal |last1=Wallach |first1=Arian D. |last2=Izhaki |first2=Ido |last3=Toms |first3=Judith D. |last4=Ripple |first4=William J. |last5=Shanas |first5=Uri |title=What is an apex predator? |journal=Oikos |volume=124 |issue=11 |year=2015 |pages=1453–1461 |doi=10.1111/oik.01977|doi-access=free |bibcode=2015Oikos.124.1453W }}</ref> They are central to the functioning of ecosystems, the regulation of disease, and the maintenance of biodiversity.<ref name="Stier 2016">{{cite journal |last1=Stier |first1=A. C. |last2=Samhouri |first2=J. F. |last3=Novak |first3=M. |last4=Marshall |first4=K. N. |last5=Ward |first5=E. J. |last6=Holt |first6=R. D. |last7=Levin |first7=P. S. |title=Ecosystem context and historical contingency in apex predator recoveries |journal=Science Advances |volume=2 |issue=5 |date=May 2016 |doi=10.1126/sciadv.1501769 |pmid=27386535 |pages=e1501769|pmc=4928970 |bibcode=2016SciA....2E1769S }}</ref> When introduced to [[subarctic]] islands, for example, [[Arctic fox]]es' predation of [[seabird]]s has been shown to turn [[grassland]] into the [[tundra]].<ref>{{cite journal |last=Croll |first=D. A. |author2=Maron, J. L. |date=March 2005 |title=Introduced predators transform subarctic islands from grassland to tundra |journal=[[Science (magazine)|Science]] |volume=307 |issue=5717 |pages=1959–1961 |doi=10.1126/science.1108485 |pmid=15790855 |display-authors=etal |bibcode=2005Sci...307.1959C |s2cid=19305264 }}</ref> Such wide-ranging effects on lower levels of an ecosystem are termed [[trophic cascade]]s. The removal of top-level predators, often through human agency, can cause or disrupt trophic cascades.<ref>{{cite web |last=Egan |first=Logan Zane |author2=Téllez, Jesús Javier |date=June 2005 |title=Effects of preferential primary consumer fishing on lower trophic level herbivores in the Line Islands |work=Stanford at Sea |publisher=[[Stanford University]] |url=http://stanford.sea.edu/research/EganTellez_Research_Project.pdf |access-date=2010-01-25 |url-status=live |archive-url=https://web.archive.org/web/20100712202445/http://stanford.sea.edu/research/EganTellez_Research_Project.pdf |archive-date=2010-07-12}}</ref><ref>{{cite journal |last=Pace |first=M. L. |author2=Cole, J. J. |display-authors=etal |date=December 1999 |title=Trophic cascades revealed in diverse ecosystems |journal=Trends in Ecology and Evolution |volume=14 |issue=12 |pages=483–488 |doi=10.1016/S0169-5347(99)01723-1 |pmid=10542455}}</ref><ref name="Berger1999">{{cite journal |last1=Berger |first1=Joel |title=Anthropogenic extinction of top carnivores and interspecific animal behaviour: implications of the rapid decoupling of a web involving wolves, bears, moose and ravens |journal=Proceedings of the Royal Society B: Biological Sciences |volume=266 |issue=1435 |year=1999 |pages=2261–2267 |doi=10.1098/rspb.1999.0917 |pmid=10629976 |url=https://sites.warnercnr.colostate.edu/joelberger/wp-content/uploads/sites/30/2017/10/1999-Ravens-Food-Web-Dynamiics-PRSB.pdf|pmc=1690453 }}</ref> For example, a reduction in the population of [[sperm whale]]s, apex predators with a [[fractional trophic level]] of 4.7, by hunting has caused an increase in the population of the large squid, with trophic level over 4 (carnivores that eat other carnivores).<ref name="Baum Worm 2009">{{cite journal |last1=Baum |first1=Julia K. |author-link1=Julia K. Baum |last2=Worm |first2=Boris |author-link2=Boris Worm |title=Cascading top-down effects of changing oceanic predator abundances |journal=Journal of Animal Ecology |volume=78 |issue=4 |year=2009 |doi=10.1111/j.1365-2656.2009.01531.x |pmid=19298616 |pages=699–714|doi-access=free |bibcode=2009JAnEc..78..699B }}</ref> This effect, called [[mesopredator release]],<ref name=Soule1988>{{cite journal |last1=Soulé |first1=Michael E. |last2=Bolger |first2=Douglas T. |last3=Alberts |first3=Allison C. |last4=Wright |first4=John |last5=Sorice |first5=Marina |last6=Hill |first6=Scott |title=Reconstructed Dynamics of Rapid Extinctions of Chaparral-Requiring Birds in Urban Habitat Islands |journal=Conservation Biology |date=March 1988 |volume=2 |issue=1 |pages=75–92 |url=https://deepblue.lib.umich.edu/bitstream/handle/2027.42/74761/j.1523-1739.1988.tb00337.x.pdf?sequence=1&isAllowed=y|doi=10.1111/j.1523-1739.1988.tb00337.x |bibcode=1988ConBi...2...75S |hdl=2027.42/74761 |hdl-access=free }}</ref> occurs in terrestrial and marine ecosystems; for instance, in North America, the ranges of all apex carnivores have contracted whereas those of 60% of mesopredators have grown in the past two centuries.<ref name="Prugh 2009">{{cite journal |last1=Prugh |first1=Laura R. |last2=Stoner |first2=Chantal J. |last3=Epps |first3=Clinton W. |last4=Bean |first4=William T. |last5=Ripple |first5=William J. |last6=Laliberte |first6=Andrea S. |last7=Brashares |first7=Justin S. |title=The Rise of the Mesopredator |journal=BioScience |volume=59 |issue=9 |year=2009 |doi=10.1525/bio.2009.59.9.9 |pages=779–791|s2cid=40484905 }}</ref>

===Conservation===
[[File:Canis lupus pack surrounding Bison.jpg|thumb|left|The [[wolf]] is both an apex predator and a [[keystone species]], affecting its prey's behaviour and the wider ecosystem.]]

Because apex predators have powerful effects on other predators, herbivores, and plants, they can be important in nature conservation.<ref name="EstesTerborgh2011">{{cite journal |last1=Estes |first1=James A. |last2=Terborgh |first2=John |last3=Brashares |first3=Justin S. |last4=Power |first4=Mary E. |last5=Berger |first5=Joel |last6=Bond |first6=William J. |last7=Carpenter |first7=Stephen R. |last8=Essington |first8=Timothy E. |last9=Holt |first9=Robert D. |last10=Jackson |first10=Jeremy B. C. |last11=Marquis |first11=Robert J. |last12=Oksanen |first12=Lauri |last13=Oksanen |first13=Tarja |last14=Paine |first14=Robert T. |last15=Pikitch |first15=Ellen K. |last16=Ripple |first16=William J. |last17=Sandin |first17=Stuart A. |last18=Scheffer |first18=Marten |last19=Schoener |first19=Thomas W. |last20=Shurin |first20=Jonathan B. |last21=Sinclair |first21=Anthony R. E. |last22=Soulé |first22=Michael E. |last23=Virtanen |first23=Risto |last24=Wardle |first24=David A. |display-authors=6 |title=Trophic Downgrading of Planet Earth |journal=Science |volume=333 |issue=6040 |year=2011 |pages=301–306 |doi=10.1126/science.1205106|pmid=21764740 |citeseerx=10.1.1.701.8043 |bibcode=2011Sci...333..301E |s2cid=7752940 }}</ref> Humans have hunted many apex predators close to extinction, but in some parts of the world, these predators are now returning.<ref name="SillimanHughes2018">{{cite journal |last1=Silliman |first1=Brian R. |last2=Hughes |first2=Brent B. |last3=Gaskins |first3=Leo C. |last4=He |first4=Qiang |last5=Tinker |first5=M. Tim |last6=Read |first6=Andrew |last7=Nifong |first7=James |last8=Stepp |first8=Rick |title=Are the ghosts of nature's past haunting ecology today? |journal=Current Biology |volume=28 |issue=9 |year=2018 |pages=R532–R537 |doi=10.1016/j.cub.2018.04.002|pmid=29738721 |doi-access=free |bibcode=2018CBio...28.R532S }}</ref> They are increasingly threatened by [[climate change]]. For example, the [[polar bear]] requires extensive areas of [[sea ice]] to hunt its prey, typically seals, but climate change is shrinking the sea ice of the Arctic, forcing polar bears to fast on land for increasingly long periods.<ref>{{cite web |url=http://pbsg.npolar.no/en/issues/threats/climate-change.html |title=Climate impacts on polar bears |publisher=Polar Bear Specialist Group |date=27 January 2009 |access-date=9 September 2018}}</ref>

Dramatic changes in the [[Greater Yellowstone Ecosystem]] were recorded after the [[gray wolf]], both an apex predator and a [[keystone species]] (one with a large effect on its ecosystem), was reintroduced to [[Yellowstone National Park]] in 1995 as a [[conservation biology|conservation]] measure. [[Elk]], the wolves' primary prey, became less abundant and changed their behavior, freeing [[riparian zone]]s from constant grazing and allowing [[willow]]s, [[aspen]]s, and [[Populus sect. Aigeiros|cottonwoods]] to flourish, creating [[habitat]]s for [[North American beaver|beaver]], [[moose]], and scores of other species.<ref>Lister, Brad &amp; McDaniel, Carl. [http://www.bioinfo.rpi.edu/~bystrc/pub/artWolves.pdf "The wolves of Yellowstone"] {{webarchive|url=https://web.archive.org/web/20110720041242/http://www.bioinfo.rpi.edu/~bystrc/pub/artWolves.pdf |date=2011-07-20 }} (2006-04-17), p. 2. Retrieved 2010-01-25.</ref> In addition to their effect on prey species, the wolves' presence also affected one of the park's [[vulnerable species]], the [[grizzly bear]]: emerging from [[hibernation]], having fasted for months, the bears chose to [[Scavenger|scavenge]] wolf kills,<ref>{{cite web |last=Levy |first=Sharon |date=November 2002 |title=Top Dogs |url=https://www.newscientist.com/article/mg17623675.300-top-dogs.html |access-date=2010-01-25 |url-status=live |archive-url=https://web.archive.org/web/20090606182407/http://www.newscientist.com/article/mg17623675.300-top-dogs.html |archive-date=2009-06-06 }}</ref> especially during the autumn as they prepared to hibernate once again.<ref name="Wilmers">{{cite web |last=Wilmers |first=Christopher C. |year=2004 |title=The gray wolf – scavenger complex in Yellowstone National Park |url=http://www.cnr.berkeley.edu/~getz/dissertation/WilmersDis.pdf |access-date=2010-01-25 |url-status=live |archive-url=https://web.archive.org/web/20100712023645/http://www.cnr.berkeley.edu/%7egetz/dissertation/WilmersDis.pdf |archive-date=2010-07-12 |pages=56, 90 and throughout}}</ref> The grizzly bear gives birth during hibernation, so the increased food supply is expected to produce an increase in the number of cubs observed.<ref name=Robbins1998>{{cite journal |last=Robbins |first=Jim |title=Weaving a new web: wolves change an ecosystem |journal=Smithsonian Zoogoer |publisher=[[Smithsonian Institution]] |date=May–June 1998 |volume=27 |issue=3 |url=http://nationalzoo.si.edu/Publications/ZooGoer/1998/3/weavingwolfweb.cfm |access-date=2010-01-25 |archive-url=https://web.archive.org/web/20100210224148/http://nationalzoo.si.edu/Publications/ZooGoer/1998/3/weavingwolfweb.cfm |archive-date=10 February 2010 |url-status=live}}</ref> Dozens of other species, including eagles, ravens, [[magpie]]s, coyotes, and [[American black bear|black bear]]s have also been documented as scavenging from wolf kills within the park.<ref name="DobsonWilmers2005">{{cite journal |last1=Dobson |first1=Andy P. |last2=Wilmers |first2=Christopher C |last3=Getz |first3=Wayne M. |title=Gray Wolves as Climate Change Buffers in Yellowstone |journal=PLOS Biology |volume=3 |issue=4 |year=2005 |pages=e92 |doi=10.1371/journal.pbio.0030092|pmid=15757363 |pmc=1064850 |doi-access=free }}</ref>{{-}}

==Human trophic level==
{{further|Trophic level}}
[[File:21 Walrus Hunt 1999.jpg|thumb|[[Human]]s sometimes live by hunting other animals for food and materials such as fur, sinew, and bone, as in this [[walrus]] hunt in the Arctic, but humans' status as apex predators is debated.]]

Ecologists have debated whether [[human]]s are apex predators. For instance, Sylvain Bonhommeau and colleagues argued in 2013 that across the global food web, a fractional human trophic level (HTL) can be calculated as the mean trophic level of every species in the human diet, weighted by the proportion that that species forms in the diet. This analysis gives an average HTL of 2.21, varying between 2.04 (for [[Burundi]], with a 96.7% plant-based diet) and 2.57 (for [[Iceland]], with 50% meat and fish, 50% plants). These values are comparable to those of non-apex predators such as the [[anchovy]] or [[pig]].<ref name="BonhommeauDubroca2013">{{cite journal |last1=Bonhommeau |first1=S. |last2=Dubroca |first2=L. |last3=Le Pape |first3=O. |last4=Barde |first4=J. |last5=Kaplan |first5=D. M. |last6=Chassot |first6=E. |last7=Nieblas |first7=A.-E. |title=Eating up the world's food web and the human trophic level |journal=Proceedings of the National Academy of Sciences |volume=110 |issue=51 |year=2013 |pages=20617–20620 |doi=10.1073/pnas.1305827110|pmid=24297882 |pmc=3870703 |bibcode=2013PNAS..11020617B |doi-access=free }}</ref>

However, Peter D. Roopnarine criticized Bonhommeau's approach in 2014, arguing that humans are apex predators and that the HTL was based on terrestrial farming where indeed humans have a low trophic level, mainly eating producers (crop plants at level 1) or primary consumers (herbivores at level 2), which as expected places humans at a level slightly above 2. Roopnarine instead calculated the position of humans in two marine ecosystems, a Caribbean coral reef and the [[Benguela current|Benguela]] system near South Africa. In these systems, humans mainly eat predatory fish and have a fractional trophic level of 4.65 and 4.5, respectively, which in Roopnarine's view makes those humans apex predators.{{efn|However, humans had a network trophic level (NTL) of 4.27 in the coral reef system, compared to an NTL of 4.8 for the [[blacktip shark]] in the same system. Therefore, humans were not the topmost apex predator there.<ref name="Roopnarine2014"/>}}<ref name="Roopnarine2014">{{cite journal |last1=Roopnarine |first1=Peter D. |title=Humans are apex predators |journal=Proceedings of the National Academy of Sciences |volume=111 |issue=9 |year=2014 |pages=E796 |doi=10.1073/pnas.1323645111 |pmid=24497513 |pmc=3948303 |bibcode=2014PNAS..111E.796R |doi-access=free }}</ref>

In 2021, Miki Ben-Dor and colleagues compared human biology to that of animals at various trophic levels. Using metrics as diverse as tool use and acidity of the stomach, they concluded that humans evolved as apex predators, diversifying their diets in response to the disappearance of most of the megafauna that had once been their primary source of food.<ref name="Ben-Dor2021">{{cite journal |last1=Ben-Dor |first1=Miki |last2=Sirtoli |first2=Raphael |last3=Barkai |first3=Ran |title=The evolution of the human trophic level during the Pleistocene |journal=[[American Journal of Physical Anthropology]] |year=2021 |volume=175 |pages=27–56 |doi=10.1002/ajpa.24247 |pmid=33675083 |doi-access=free }}</ref>
<!-- please think twice (ok, three times, and go for a cold shower first) before adding anything else here, it's getting excessive for a general article. Pr [[WP:COATRACK]] (or just [[WP:DUE]]) we'll soon need to split it off to a separate article and just summarize it here very briefly -->

==Evolutionary history==
[[File:20191203 Anomalocaris canadensis.png|thumb|left|''[[Anomalocaris]]'' was an apex predator in the [[Cambrian]] seas.<ref name=Callaway2011/>]]

{{Further|Predation#Evolutionary history}}

Apex predators are thought to have existed since at least the [[Cambrian]] period, around 500 million years ago. Extinct species cannot be directly determined to be apex predators as their behavior cannot be observed, and clues to ecological relationships, such as bite marks on bones or shells, do not form a complete picture. However, indirect evidence such as the absence of any discernible predator in an environment is suggestive. ''[[Anomalocaris]]'' was an aquatic apex predator, in the Cambrian. Its mouthparts are clearly predatory, and there were no larger animals in the seas at that time.<ref name=Callaway2011>{{cite journal |last=Callaway |first=Ewen |title=Cambrian super-predators grew large in arms race |journal=Nature |date=2011 |doi=10.1038/news.2011.318}}</ref>

Carnivorous [[theropod]] dinosaurs including ''[[Allosaurus]]''<ref name="ZannoMakovicky2013">{{cite journal |last1=Zanno |first1=Lindsay E. |last2=Makovicky |first2=Peter J. |title=Neovenatorid theropods are apex predators in the Late Cretaceous of North America |journal=Nature Communications |volume=4 |issue=1 |pages=2827 |date=22 November 2013 |doi=10.1038/ncomms3827|pmid=24264527 |bibcode=2013NatCo...4.2827Z |doi-access=free }}</ref> and ''[[Tyrannosaurus]]''<ref name=ValkenburghMolnar2002>{{cite journal |last1=Van Valkenburgh |first1=Blaire |last2=Molnar |first2=Ralph E. |title=Dinosaurian and Mammalian Predators Compared |journal=Paleobiology |date=2002 |volume=28 |issue=4 |pages=527–543 |jstor=3595499|doi=10.1666/0094-8373(2002)028<0527:DAMPC>2.0.CO;2 |bibcode=2002Pbio...28..527V |s2cid=85725299 |url=https://zenodo.org/record/890154 }}</ref> are theorized to have been apex predators, based on their size, morphology, and dietary needs.

A [[Permian]] shark, ''[[Triodus|Triodus sessilis]]'', was discovered containing two [[amphibians]] (''[[Archegosaurus decheni]]'' and ''[[Cheliderpeton latirostre]]''), one of which had consumed a fish, ''[[Acanthodes|Acanthodes bronni]]'', showing that the shark had lived at a trophic level of at least 4.{{efn|Its [[trophic level]] would be exactly 4 if the fish's prey were pure herbivores, higher if the prey were themselves carnivorous.}}<ref name="KriwetWitzmannKlugHeidtke2008">{{cite journal |last1=Kriwet |first1=J. |last2=Witzmann |first2=F. |last3=Klug |first3=S. |last4=Heidtke |first4=U. H.J |title=First direct evidence of a vertebrate three-level trophic chain in the fossil record |journal=Proceedings of the Royal Society B: Biological Sciences |volume=275 |issue=1631 |date=2008 |doi=10.1098/rspb.2007.1170 |pmid=17971323 |pmc=2596183 |pages=181–186}}</ref>

Among more recent fossils, the saber-tooth cats, like ''[[Smilodon]]'', are considered to have been apex predators in the [[Cenozoic]].<ref name=Werdelin2018>{{cite book |last1=Werdelin |first1=Lars |last2=McDonald |first2=H. G. |last3=Shaw |first3=Christopher A. |title=Smilodon: The Iconic Sabertooth |url=https://books.google.com/books?id=XnJVDwAAQBAJ&pg=PA204 |year=2018 |publisher=Johns Hopkins University Press |isbn=978-1-4214-2556-6 |page=204}}</ref>

==Interactions with humans==
[[File:37-svaghi, caccia,Taccuino Sanitatis, Casanatense 4182..jpg|thumb|upright|Dogs have been used in hunting for many millennia, as in this 14th century French depiction of a [[Wild boar|boar]] hunt.]]

===Hunting===
{{Further|Hunting}}

Humans hunted with apex predators in the form of wolves, and in turn with [[domestic dog]]s, for 40,000 years; this collaboration may have helped modern humans to outcompete the [[Neanderthals]].<ref>{{cite news |author=McKie, Robin |url=https://www.theguardian.com/science/2015/mar/01/hunting-with-wolves-humans-conquered-the-world-neanderthal-evolution |title=How hunting with wolves helped humans outsmart the Neanderthals |work=[[The Guardian]] |date=1 March 2015 |access-date=14 October 2018}}</ref><ref name="Shipman 2015">{{cite book |last=Shipman |first=Pat |title=The invaders : how humans and their dogs drove Neanderthals to extinction |publisher=Harvard University Press |year=2015 |isbn=978-0-674-73676-4 |oclc=893897294}}</ref> Humans still hunt [[hunting dog|with dogs]], which have often been bred as [[gun dog]]s to [[Pointing breed|point to]], [[Flushing dog|flush out]], or [[Retriever|retrieve prey]].<ref>{{cite news |url=https://www.telegraph.co.uk/pets/essentials/seven-types-of-dog/ |archive-url=https://ghostarchive.org/archive/20220111/https://www.telegraph.co.uk/pets/essentials/seven-types-of-dog/ |archive-date=2022-01-11 |url-access=subscription |url-status=live |title=The 7 categories of dog |date=10 March 2017 |publisher=[[The Daily Telegraph]]}}{{cbignore}}</ref> The [[Portuguese Water Dog]] was used to drive fish into nets.<ref>{{cite web |title=Portuguese Water Dog |url=https://www.thekennelclub.org.uk/services/public/breed/pdf.ashx?breed=5135&item=1111011 |publisher=[[The Kennel Club]] |access-date=14 October 2018 |archive-date=14 October 2018 |archive-url=https://web.archive.org/web/20181014130131/https://www.thekennelclub.org.uk/services/public/breed/pdf.ashx?breed=5135&item=1111011 |url-status=dead }}</ref> Several breeds of dog have been used to [[Scottish Deerhound|chase large prey such as deer]] and wolves.<ref>{{cite book |author=Fergus, Charles |title=Gun Dog Breeds, A Guide to Spaniels, Retrievers, and Pointing Dogs |publisher=The Lyons Press |date=2002 |isbn=978-1-58574-618-7}}</ref>

[[Eagle]]s and [[falcon]]s, which are apex predators, are used in [[falconry]], hunting birds or mammals.<ref>{{cite web |title=History of Falconry |url=http://www.thefalconrycentre.co.uk/bird-info/conservation/nocturnal-raptors/history-falconry/ |publisher=The Falconry Centre |access-date=22 April 2016 |archive-date=29 May 2016 |archive-url=https://web.archive.org/web/20160529023926/http://thefalconrycentre.co.uk/bird-info/conservation/nocturnal-raptors/history-falconry/ |url-status=dead }}</ref> Tethered [[cormorant]]s, also top predators,<ref>{{cite journal |last1=Rudstam |first1=Lars G. |last2=VanDeValk |first2=Anthony J. |title=Cormorant Predation and the Population Dynamics of Walleye and Yellow Perch in Oneida Lake |journal=Ecological Applications |date=February 2004 |volume=14 |issue=1 |pages=149–163 |jstor=4493527 |quote=providing the opportunity to study the effects of an increase of a top predator on an existing predator-prey system|doi=10.1890/03-5010 |bibcode=2004EcoAp..14..149R }}</ref> have been [[Cormorant fishing|used to catch fish]].<ref name="King2013">{{cite book |author=King, Richard J. |title=The Devil's Cormorant: A Natural History |url=https://books.google.com/books?id=ucGyAAAAQBAJ&pg=PA9 |date=2013 |publisher=University of New Hampshire Press |isbn=978-1-61168-225-0 |page=9}}</ref>

===Ecotourism===
{{Further|Ecotourism}}
[[File:Tiger shark size.svg|thumb|left|[[Tiger shark]]s are popular [[ecotourism]] subjects, but their ecosystems may be affected by the food provided to attract them.]]

[[Ecotourism]] sometimes relies on apex predators to attract business.<ref name="HammerschlagGallagher2012"/><ref name="Macdonald 2017"/> Tour operators may in consequence decide to intervene in ecosystems, for example by providing food to attract predators to areas that can conveniently be visited.<ref name="HammerschlagGallagher2012"/> This in turn can have effects on predator population and therefore on the wider ecosystem.<ref name="HammerschlagGallagher2012"/> As a result, provisioning of species such as the [[tiger shark]] is controversial, but its effects are not well established by empirical evidence.<ref name="HammerschlagGallagher2012">{{cite journal |last1=Hammerschlag |first1=Neil |last2=Gallagher |first2=Austin J. |last3=Wester |first3=Julia |last4=Luo |first4=Jiangang |last5=Ault |first5=Jerald S. |title=Don't bite the hand that feeds: assessing ecological impacts of provisioning ecotourism on an apex marine predator |journal=Functional Ecology |volume=26 |issue=3 |year=2012 |pages=567–576 |doi=10.1111/j.1365-2435.2012.01973.x|doi-access=free |bibcode=2012FuEco..26..567H }}</ref> Other affected apex predators include [[big cats]] and crocodiles.<ref name="Macdonald 2017">{{cite journal |last1=Macdonald |first1=Catherine |last2=Gallagher |first2=Austin J. |last3=Barnett |first3=Adam |last4=Brunnschweiler |first4=Juerg |last5=Shiffman |first5=David S. |last6=Hammerschlag |first6=Neil |title=Conservation potential of apex predator tourism |journal=Biological Conservation |volume=215 |year=2017 |doi=10.1016/j.biocon.2017.07.013 |pages=132–141|bibcode=2017BCons.215..132M }}</ref>{{-}}

===Rewilding===
{{Further|Rewilding (conservation biology)}}
[[File:Lynx Lynx (10597310204).jpg|thumb|The reintroduction of predators like the [[Eurasian lynx|lynx]] is attractive to conservationists, but alarming to farmers.]]

In some densely populated areas like the [[British Isles]], all the large native predators like the wolf, bear, wolverine and lynx have become [[Extirpate|extirpated]], allowing herbivores such as deer to multiply unchecked except by hunting.<ref name="Jones2015">{{cite web |last1=Jones |first1=Lucy |title=The rewilding plan that would return Britain to nature |url=https://www.bbc.co.uk/earth/story/20150604-can-we-make-britain-wild-again |publisher=BBC |access-date=6 June 2018 |quote=wolves, bears and lynx roamed the land. ... Humans chopped down the trees to make space for farms, and hunted the large animals to extinction, leaving plant-eaters to decimate the country's flora. Britain is now one of the few countries in the world that doesn't have top predators.}}</ref> In 2015, plans were made to reintroduce [[lynx]] to the counties of [[Norfolk]], [[Cumbria]], and [[Northumberland]] in England, and [[Aberdeenshire]] in Scotland as part of the [[Rewilding (conservation biology)|rewilding]] movement.<ref name=Lister2015/> The reintroduction of large predators is controversial, in part because of concern among farmers for their [[livestock]].<ref name=Lister2015/> Conservationists such as Paul Lister propose instead to allow wolves and bears to hunt their prey in a "managed environment" on large fenced reserves; however, this undermines the objective of rewilding.<ref name=Lister2015>{{cite news |last1=Lister |first1=Paul |title=Bring on a few more apex predators |url=https://www.telegraph.co.uk/news/earth/environment/conservation/11566555/Bring-on-a-few-more-apex-predators.html |archive-url=https://ghostarchive.org/archive/20220111/https://www.telegraph.co.uk/news/earth/environment/conservation/11566555/Bring-on-a-few-more-apex-predators.html |archive-date=2022-01-11 |url-access=subscription |url-status=live |access-date=14 March 2018 |work=The Daily Telegraph |date=28 April 2015}}{{cbignore}}</ref>

==Notes==
{{Notelist}}

==References==
{{Reflist|30em}}

==External links==
* [https://www.ibiology.org/ecology/apex-predators/ The Ecological Function of Apex Predators]: talk by Prof. James Estes (UC Santa Cruz)

{{Feeding}}
{{Modelling ecosystems}}
{{Authority control}}

{{DEFAULTSORT:Apex Predator}}
[[Category:Predation]]
[[Category:Apex predators|*]]