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====Fossil record==== {{See also|Evolution of the wolf#Fossil record}} [[Xiaoming Wang (paleontologist)|Xiaoming Wang]] and [[Richard H. Tedford]], one of the foremost [[Author citation (zoology)|authorities]] on carnivore evolution,<ref>{{cite magazine|title=Natural History: Canid Family Ties |year=2008|magazine=The Magazine of the American Museum of Natural History|url=https://archive.org/stream/naturalhistory11706unse#page/22|page=22|volume=117|number=6|publisher=American Museum of Natural History|location=New York}}</ref> proposed that the genus ''Canis'' was the descendant of the coyote-like ''[[Eucyon davisi]]'' and its remains first appeared in the [[Miocene]] 6{{nbsp}}million years ago (Mya) in the southwestern US and Mexico. By the [[Pliocene]] (5{{nbsp}}Mya), the larger ''[[Canis lepophagus]]''<ref name=fossilworks1/> appeared in the same region and by the [[early Pleistocene]] (1{{nbsp}}Mya) ''C.{{nbsp}}latrans'' (the coyote) was in existence. They proposed that the progression from ''Eucyon davisi'' to ''C.{{nbsp}}lepophagus'' to the coyote was linear evolution.{{sfn|Wang|Tedford|2008|p=58}} ''C.{{nbsp}}latrans'' and ''C.{{nbsp}}aureus'' are closely related to ''[[Canis edwardii|C.{{nbsp}}edwardii]]'', a species that appeared earliest spanning the mid-[[Blancan]] ([[late Pliocene]]) to the close of the [[Irvingtonian]] (late Pleistocene), and coyote remains indistinguishable from ''C. latrans'' were contemporaneous with ''C.{{nbsp}}edwardii'' in North America.{{sfn|Tedford|Wang|Taylor|2009|pp=175, 180}} Johnston describes ''C.{{nbsp}}lepophagus'' as having a more slender skull and skeleton than the modern coyote.{{sfn|Johnston|1938|p=385}} Ronald Nowak found that the early populations had small, delicate, narrowly proportioned skulls that resemble small coyotes and appear to be ancestral to ''C. latrans''.{{sfn|Nowak|2003|p=241}} ''C. lepophagus'' was similar in weight to modern coyotes, but had shorter limb bones that indicate a less [[cursorial]] lifestyle. The coyote represents a more primitive form of ''Canis'' than the gray wolf, as shown by its relatively small size and its comparatively narrow skull and jaws, which lack the grasping power necessary to hold the large prey in which wolves specialize. This is further corroborated by the coyote's [[sagittal crest]], which is low or totally flattened, thus indicating a weaker bite than the wolves. The coyote, unlike the wolf, is not a specialized carnivore, as shown by the larger chewing surfaces on the [[molar (tooth)|molars]], reflecting the species' relative dependence on vegetable matter. In these respects, the coyote resembles the fox-like progenitors of the genus more so than the wolf.<ref name=nowak1978/> The oldest fossils that fall within the range of the modern coyote date to 0.74β0.85 [[Megaannum|Ma]] (million years) in Hamilton Cave, West Virginia; 0.73 Ma in Irvington, California; 0.35β0.48 Ma in Porcupine Cave, Colorado, and in Cumberland Cave, Pennsylvania.{{sfn|Tedford|Wang|Taylor|2009|p=136}} Modern coyotes arose 1,000 years after the [[Quaternary extinction event]].<ref name="meachen2012"/> Compared to their modern [[Holocene]] counterparts, [[Pleistocene coyote]]s (''C.{{nbsp}}l. orcutti'') were larger and more robust, likely in response to larger [[Competition (biology)|competitors]] and prey.<ref name="meachen2012">{{cite journal | doi = 10.1073/pnas.1113788109|pmid=22371581| title = Evolution in coyotes (''Canis latrans'') in response to the megafaunal extinctions| journal = Proceedings of the National Academy of Sciences| volume = 109| issue = 11| pages = 4191β6| year = 2012| last1 = Meachen | first1 = J. A. | last2 = Samuels | first2 = J. X. |issn=1091-6490|oclc=475396714|bibcode=2012PNAS..109.4191M | pmc=3306717| url=http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1043&context=natlpark|doi-access=free}}</ref> Pleistocene coyotes were likely more specialized carnivores than their descendants, as their teeth were more adapted to shearing meat, showing fewer grinding surfaces suited for processing vegetation.<ref name="meachen2014">{{Cite journal | doi = 10.1371/journal.pone.0116041|pmid=25551387| title = Ecological Changes in Coyotes (''Canis latrans'') in Response to the Ice Age Megafaunal Extinctions| journal = PLOS ONE| volume = 9| issue = 12| page = e116041| year = 2014| last1 = Meachen | first1 = J. A. | last2 = Janowicz | first2 = A. C. | last3 = Avery | first3 = J. E. | last4 = Sadleir | first4 = R. W. |bibcode = 2014PLoSO...9k6041M | pmc=4281224|doi-access=free}}</ref> Their reduction in size occurred within 1,000 years of the Quaternary extinction event, when their large prey died out.<ref name="meachen2012"/> Furthermore, Pleistocene coyotes were unable to exploit the big-game hunting [[Ecological niche|niche]] left vacant after the extinction of the [[dire wolf]] (''Aenocyon{{nbsp}}dirus''), as it was rapidly filled by gray wolves, which likely actively killed off the large coyotes, with [[natural selection]] favoring the modern gracile morph.<ref name="meachen2014"/>
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