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The Pleistocene coyote (''Canis latrans orcutti''), also known as the Ice Age coyote, is an extinct subspecies of
coyote The coyote (''Canis latrans'') is a species of canine native to North America. It is smaller than its close relative, the wolf, and slightly smaller than the closely related eastern wolf and red wolf. It fills much of the same ecological nich ...
that lived in western
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
during the
Late Pleistocene The Late Pleistocene is an unofficial age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a stratigraphic perspective. It is intended to be the fourth division of the Pleistocene Epoch withi ...
era. Most remains of the subspecies were found in southern
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
, though at least one was discovered in
Idaho Idaho ( ) is a U.S. state, state in the Pacific Northwest region of the Western United States. To the north, it shares a small portion of the Canada–United States border with the province of British Columbia. It borders the states of Monta ...
. It was part of a North American carnivore guild that included other
canid Canidae (; from Latin, '' canis'', " dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found withi ...
s like foxes,
gray wolves The wolf (''Canis lupus''; : wolves), also known as the gray wolf or grey wolf, is a large canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, and gray wolves, as popularly u ...
, and
dire wolves The dire wolf (''Aenocyon dirus'' ) is an extinct canine. It is one of the most famous prehistoric carnivores in North America, along with its extinct competitor ''Smilodon''. The dire wolf lived in the Americas and eastern Asia during the Lat ...
.Meachen, J., Samuels, J. (2012)
Evolution in coyotes (Canis latrans) in response to the megafaunal extinctions
''PNAS'' : 10.1073/pnas.1113788109
Some studies suggest that the Pleistocene "coyote" was not in fact a coyote, but rather an extinct western population of the
red wolf The red wolf (''Canis rufus'') is a canine native to the southeastern United States. Its size is intermediate between the coyote (''Canis latrans'') and gray wolf (''Canis lupus''). The red wolf's taxonomic classification as being a separate s ...
(''C. rufus'').


Description

Compared to their modern Holocene counterparts, Pleistocene coyotes were larger and more robust, weighing , likely in response to larger competitors and prey rather than
Bergmann's rule Bergmann's rule is an ecogeographical rule that states that within a broadly distributed taxonomic clade, populations and species of larger size are found in colder environments, while populations and species of smaller size are found in warmer r ...
. Their skulls and jaws were significantly thicker and deeper than in modern coyotes, with a shorter and broader rostrum and wider
carnassial Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified f ...
(denoting the large upper premolar and lower molar teeth of a carnivore, adapted for shearing flesh) teeth. These adaptions allowed it to cope with higher levels of stress, when it killed larger prey, compared to modern coyotes. Pleistocene coyotes were also likely more specialized carnivores than their descendants, as their teeth were more adapted to shearing meat, showing fewer grinding surfaces which were better suited for processing vegetation. The lower jaw was also deeper, and the molars showed more signs of wear and breakage than modern populations, thus indicating that the animals consumed more bone than today.Meachen JA, Janowicz AC, Avery JE, Sadleir RW (2014
Ecological Changes in Coyotes (''Canis latrans'') in Response to the Ice Age Megafaunal Extinctions
''PLoS ONE'' 9(12): e116041. doi:10.1371/journal.pone.0116041
Behaviorally, it is likely to have been more social than the modern coyote, as its remains are the third most common in the
La Brea Tar Pits La Brea Tar Pits is an active paleontological research site in urban Los Angeles. Hancock Park was formed around a group of tar pits where natural asphalt (also called asphaltum, bitumen, or pitch; ''brea'' in Spanish) has seeped up from the gr ...
, after dire wolves and
sabre-toothed cat Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae (true cats). They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million unt ...
s, both thought to be gregarious species. Their reduction in size occurred within 1,000 years of the occurrence of the
Quaternary extinction event The Quaternary period (from 2.588 ± 0.005 million years ago to the present) has seen the extinctions of numerous predominantly megafaunal species, which have resulted in a collapse in faunal density and diversity and the extinction of key ecolog ...
, when the climate changed and the majority of their larger prey became extinct. Furthermore, Pleistocene coyotes were unable to successfully exploit the big game hunting
niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development * Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...
left vacant after the extinction of the dire wolf, as that gap was rapidly filled by gray wolves. These gray wolves are likely to have actively killed off the larger-bodied coyotes, with
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
favoring the modern gracile morph.
Human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
predation on the Pleistocene coyote's dwindling prey base may have also impacted the animal's change in morphology.


''Canis latrans harriscrooki''

''Canis latrans harriscrooki'' (Slaughter, 1961)Slaughter, B.H. (1961) A new coyote in the Late Pleistocene of Texas. Journal of Mammalogy 42(4):503–509. is another extinct Late Pleistocene coyote that once inhabited what is now Texas. Slaughter described it as being wolf-like and was distinguished from other coyotes by a well-developed posterior cusp on its p2 (the second premolar on its mandible), a longer tooth row relative to the depth of its mandible, a reduced distance between premolars, and a more vertical descending ramus. The cusp dentition was also found in two specimens from Mexico and one from Honduras. Slaughter identified some affinity with ''C. l. hondurensis''. Nowak proposed the hypothesis that a warm-adapted coyote that was more wolf-like than modern coyotes once inhabited Pleistocene Texas and might still be represented by ''C.l. hondurensis''.


''Canis latrans riviveronis''

''Canis riviveronis'' (Hay, 1917)O. P. Hay. 1917. Vertebrata mostly from Stratum No. 3, at Vero, Florida, together with descriptions of new species. Florida State Geological Survey Annual Report 9:43-68
/ref> is a coyote that once lived in Florida during the Pleistocene. The specimen is described as a coyote, but it being ''latrans'' was questionable. It differed from the extant coyote by having the anterior lobe of the carnassial relatively shorter, and the teeth broader. It was not a wolf nor an Indian dog.


Descendants

In 2021, a
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
analysis of modern and extinct North American wolf-like canines indicates that the
Beringian wolf The Beringian wolf is an extinct population of wolf (''Canis lupus'') that lived during the Ice Age. It inhabited what is now modern-day Alaska, Yukon, and northern British Columbia. Some of these wolves survived well into the Holocene. The ...
was the ancestor of the southern wolf clade, which includes the Mexican wolf and the extinct Great Plains wolf. The Mexican wolf is the most ancestral of the gray wolves that live in North America today. The modern coyote appeared around 10,000 years ago. The most genetically basal coyote mDNA clade pre-dates the Late Glacial Maximum and is a haplotype that can only be found in the Eastern wolf. This implies that the large, wolf-like Pleistocene coyote was the ancestor of the Eastern wolf. Further, another ancient haplotype detected in the Eastern wolf can be found only in the Mexican wolf. The study proposes that Pleistocene coyote and Beringian wolf admixture led to the Eastern wolf long before the arrival of the modern coyote and the modern wolf. While not necessarily direct descendants, some populations of modern
Eastern coyote An eastern coyote in the snow near the West Virginia–Virginia state line">Virginia.html" ;"title="West Virginia–Virginia">West Virginia–Virginia state line The eastern coyote is a wild North American canine hybrid with both coyote and wolf ...
which originated through a combination of hybridization with Eastern wolf and other canine populations as well as selective pressures favoring a larger body size to exploit a niche left vacant by the local extinction of gray wolves in the eastern United States, have adapted a form superficially similar to the Pleistocene coyote. Similarities include the enlarged
Sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exception ...
for larger jaw muscles, more robust teeth, a higher tendency of pack living and hunting behavior, and a larger body size compared to western coyote populations. In some areas, eastern coyotes regularly reach the same size as their Pleistocene counterparts."Eastern Coyote Wildlife Note"
/ref>


Debates over identity

In 2021, another mitochondrial DNA analysis of Pleistocene coyote DNA and historic red/eastern wolf material found that the Pleistocene coyote had an independent origin from mid-continent coyotes, and also found evidence of historic
red wolf The red wolf (''Canis rufus'') is a canine native to the southeastern United States. Its size is intermediate between the coyote (''Canis latrans'') and gray wolf (''Canis lupus''). The red wolf's taxonomic classification as being a separate s ...
and eastern wolf populations carrying
haplotype A haplotype ( haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material ( DNA) which is inherited from two parents. Normally these organisms have their DNA o ...
s from Pleistocene coyotes, but found no evidence of these haplotypes in mid-continent coyotes. For this reason, the study postulated that the Pleistocene "coyote" and the common ancestor of red and eastern wolves had connectivity between 50,000 to 60,000 years ago. This relatively recent connectivity was interpreted as indicating conspecificity, leading to the conclusion that the Pleistocene "coyote" was actually an extinct western population of the red wolf. The study postulated that a prehistoric expansion of true coyotes into California led to the outbreeding and extirpation of the western red wolf.


References

{{Taxonbar, from=Q20720501 Coyotes Coyote, Pleistocene Coyote, Pleistocene Coyote, Pleistocene Coyote, Pleistocene Late Pleistocene Coyote, Pleistocene Coyote, Pleistocene Extinct animals of the United States Fauna of California Natural history of Los Angeles County, California Pleistocene California Fossil taxa described in 1910