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== Evolutionary history == {{further|History of life}} Predation dates from before the rise of commonly recognized carnivores by hundreds of millions (perhaps billions) of years. Predation has evolved repeatedly in different groups of organisms.<ref name="Bengtson2002"/><ref>{{cite journal |author=Abrams, P. A. |date=2000 |title=The evolution of predator-prey interactions: theory and evidence |journal=Annual Review of Ecology and Systematics |volume=31 |issue=1 |pages=79–105|doi=10.1146/annurev.ecolsys.31.1.79 |bibcode=2000AnRES..31...79A }}</ref> The rise of [[eukaryotic]] cells at around 2.7 Gya, the rise of multicellular organisms at about 2 Gya, and the rise of mobile predators (around 600 Mya - 2 Gya, probably around 1 Gya) have all been attributed to early predatory behavior, and many very early remains show evidence of boreholes or other markings attributed to small predator species.<ref name=Bengtson2002 /> It likely triggered major evolutionary transitions including the arrival of [[Cell (biology)|cell]]s, [[eukaryote]]s, [[sexual reproduction]], [[multicellularity]], increased size, mobility (including [[insect flight]]<ref name=GrimaldiEngel2005/>) and armoured shells and exoskeletons.<ref name="Bengtson2002"/> The earliest predators were microbial organisms, which engulfed or grazed on others. Because the fossil record is poor, these first predators could date back anywhere between 1 and over 2.7 Gya (billion years ago).<ref name=Bengtson2002/> Predation visibly became important shortly before the [[Cambrian]] period—around {{Ma |550}}—as evidenced by the almost simultaneous development of [[calcification]] in animals and algae,<ref name="Grant1991">{{cite journal |author1=Grant, S. W. F. |author2=Knoll, A. H. |author3=Germs, G. J. B. |year=1991 |title=Probable Calcified Metaphytes in the Latest Proterozoic Nama Group, Namibia: Origin, Diagenesis, and Implications |journal=Journal of Paleontology |volume=65 |issue=1 |pages=1–18 |pmid=11538648 |jstor=1305691|doi=10.1017/S002233600002014X |bibcode=1991JPal...65....1G |s2cid=26792772 }}</ref> and predation-avoiding [[burrow]]ing. However, predators had been grazing on micro-organisms since at least {{Ma |1000}},<ref name="Bengtson2002">{{cite book |author=Bengtson, S. |year=2002 |contribution=Origins and early evolution of predation |title=The fossil record of predation. The Paleontological Society Papers 8 |editor-last1=Kowalewski |editor-first1=M. |editor-last2=Kelley |editor-first2=P. H. |pages=289–317 |publisher=The Paleontological Society |url=http://www.nrm.se/download/18.4e32c81078a8d9249800021552/Bengtson2002predation.pdf |access-date=23 July 2008 |archive-date=10 September 2008 |archive-url=https://web.archive.org/web/20080910205539/http://www.nrm.se/download/18.4e32c81078a8d9249800021552/Bengtson2002predation.pdf |url-status=dead }}</ref><ref name="Awramik1971">{{cite journal |author=Awramik, S. M. |title=Precambrian columnar stromatolite diversity: Reflection of metazoan appearance |journal=Science |volume=174 |issue=4011 |pages=825–827 |date=19 November 1971 |doi=10.1126/science.174.4011.825 |pmid=17759393 |bibcode=1971Sci...174..825A|s2cid=2302113 }}</ref><ref name="Stanley2008">{{cite journal |title=Predation defeats competition on the seafloor |author=Stanley, Steven M. |year=2008 |journal=Paleobiology |volume=34 |pages=1–21 |doi=10.1666/07026.1 |issue=1|bibcode=2008Pbio...34....1S |s2cid=83713101 }}</ref> with evidence of selective (rather than random) predation from a similar time.<ref>{{cite journal |doi=10.1042/ETLS20170153 |title=Implications of selective predation on the macroevolution of eukaryotes: Evidence from Arctic Canada |journal=Emerging Topics in Life Sciences|volume=2 |issue=2 |pages=247–255 |year=2018 |last1=Loron |first1=Corentin C. |last2=Rainbird |first2=Robert H. |last3=Turner |first3=Elizabeth C. |last4=Wilder Greenman |first4=J. |last5=Javaux |first5=Emmanuelle J.|pmid=32412621 |s2cid=92505644 }}</ref> ''[[Auroralumina attenboroughii]]'' is an Ediacaran crown-group [[cnidaria]]n (557–562 mya, some 20 million years before the Cambrian explosion) from [[Charnwood Forest]], England. It is thought to be one of the earliest predatory animals, catching small prey with its [[nematocyst]]s as modern cnidarians do.<ref name="Dunn Kenchington Parry Clark 2022">{{cite journal |last1=Dunn |first1=F. S. |last2=Kenchington |first2=C. G. |last3=Parry |first3=L. A. |last4=Clark |first4=J. W. |last5=Kendall |first5=R. S. |last6=Wilby |first6=P. R. |title=A crown-group cnidarian from the Ediacaran of Charnwood Forest, UK |journal=Nature Ecology & Evolution |date=25 July 2022 |volume=6 |issue=8 |pages=1095–1104 |doi=10.1038/s41559-022-01807-x |pmid=35879540 |pmc=9349040 |bibcode=2022NatEE...6.1095D }}</ref> The [[fossil record]] demonstrates a long history of interactions between predators and their prey from the Cambrian period onwards, showing for example that some predators drilled through the shells of [[bivalve]] and [[gastropod]] molluscs, while others ate these organisms by breaking their shells.<ref name="Kelley">{{cite book |last=Kelley |first=Patricia |title=Predator—Prey Interactions in the Fossil Record |publisher=Springer |year=2003 | isbn=978-1-4615-0161-9 | oclc=840283264 |pages=113–139, 141–176 and passim}}</ref> Among the Cambrian predators were invertebrates like the [[anomalocaridid]]s with appendages suitable for grabbing prey, large compound eyes and jaws made of a hard material like that in the [[exoskeleton]] of an insect.<ref>{{cite journal |last=Daley |first=Allison C. |title=Anomalocaridids |journal=Current Biology |volume=23 |issue=19 |pages=R860–R861 |doi=10.1016/j.cub.2013.07.008|pmid=24112975 |year=2013 |doi-access=free |bibcode=2013CBio...23.R860D }}</ref> Some of the first [[jawed fish|fish to have jaws]] were the armoured and mainly predatory [[placoderms]] of the [[Silurian]] to [[Devonian]] periods, one of which, the {{convert|6|m|abbr=on}} ''[[Dunkleosteus]]'', is considered the world's first [[vertebrate]] "superpredator", preying upon other predators.<ref name=Anderson&Westneat2009>{{cite journal |last=Anderson |first=P. S. L. |author2=Westneat, M. |title=A biomechanical model of feeding kinematics for Dunkleosteus terrelli (Arthrodira, Placodermi) |journal=Paleobiology |date=2009 |volume=35 |issue=2 |pages=251–269|doi=10.1666/08011.1|bibcode=2009Pbio...35..251A |s2cid=86203770 }}</ref><ref name="Carr2010">{{cite journal |last1=Carr |first1=Robert K. |title=Paleoecology of Dunkleosteus Terrelli (Placodermi: Arthrodira) |journal=Kirtlandia |date=2010 |volume=57 |pages=36–45 |url=https://biostor.org/reference/193162 }}</ref> [[Insect]]s developed the ability to fly in the Early [[Carboniferous]] or Late Devonian, enabling them among other things to escape from predators.<ref name=GrimaldiEngel2005>{{cite book |last1=Grimaldi |first1=David |author-link1=David Grimaldi (entomologist) |last2=Engel |first2=Michael S. |author-link2=Michael S. Engel |year=2005 |title=Evolution of the Insects |url=https://archive.org/details/evolutioninsects00grim_110 |url-access=limited |publisher=Cambridge University Press |isbn=978-0-521-82149-0 |pages=[https://archive.org/details/evolutioninsects00grim_110/page/n169 155]–160}}</ref> Among the largest predators that have ever lived were the [[theropod dinosaur]]s such as ''[[Tyrannosaurus]]'' from the [[Cretaceous]] period. They preyed upon herbivorous dinosaurs such as [[hadrosaur]]s, [[ceratopsia]]ns and [[ankylosaur]]s.<ref>{{cite journal |last1=Sampson |first1=Scott D. |last2=Loewen |first2=Mark A. |title=Tyrannosaurus rex from the Upper Cretaceous (Maastrichtian) North Horn Formation of Utah: biogeographic and paleoecologic implications |journal=Journal of Vertebrate Paleontology |date=27 June 2005 |volume=25 |issue=2 |pages=469–472 |doi=10.1671/0272-4634(2005)025[0469:TRFTUC]2.0.CO;2 |s2cid=131583311 }}</ref> <gallery class="center" mode="nolines" widths="210px" heights="180px"> File:Auroralumina attenboroughii reconstruction.jpg|''[[Auroralumina attenboroughii]]'', an [[Ediacaran]] predator (c. 560 mya). It was a stem-group [[cnidarian]], catching prey with its [[nematocyst]]s.<ref name="Dunn Kenchington Parry Clark 2022"/> File:Cambrian substrate revolution 02.png|The [[Cambrian substrate revolution]] saw life on the [[sea floor]] change from minimal burrowing (left) to a diverse burrowing fauna (right), probably to avoid new [[Cambrian]] predators. File:20191021 Peytoia nathorsti Laggania cambria.png|The [[anomalocaridid]] ''[[Peytoia]]'', a [[Cambrian]] invertebrate, probably an apex predator File:Meganeura monyi au Museum de Toulouse.jpg|''[[Meganeura monyi]]'', a predatory [[Carboniferous]] [[insect]] related to [[Dragonfly|dragonflies]], could fly to escape terrestrial predators. Its large size, with a wingspan of {{convert|65|cm|in|-1|abbr=on}}, may reflect the lack of vertebrate aerial predators at that time. File:Tyrannosaurus rex Reconstruction by Nobu Tamura.jpg|''[[Tyrannosaurus]]'', a large [[theropod dinosaur]] of the [[Cretaceous]], reconstruction File:Dunkleosteus terrelli (2024).png|[[Dunkleosteus|Dunkleosteus terrelli]], a [[Devonian]] [[placoderm]], perhaps the world's first vertebrate [[Super-predator|superpredator]], reconstruction </gallery>
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