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    Dire wolf - Wikipedia

    The dire wolf is an extinct canine. The dire wolf lived in the Americas during the Late Pleistocene and Early Holocene epochs (125,000–9,500 years ago). A putative, controversial fossil was recently reported from northeast China, but other researchers questioned the taxonomic attribution of this specimen. The species was named in 1858, four years after the first specimen had been found. Two subspeciesare recognized: Aenocyon dirus guild…

    The dire wolf is an extinct canine. The dire wolf lived in the Americas during the Late Pleistocene and Early Holocene epochs (125,000–9,500 years ago). A putative, controversial fossil was recently reported from northeast China, but other researchers questioned the taxonomic attribution of this specimen. The species was named in 1858, four years after the first specimen had been found. Two subspecies are recognized: Aenocyon dirus guildayi and Aenocyon dirus dirus. The largest collection of its fossils has been obtained from the Rancho La Brea Tar Pits in Los Angeles.

    Dire wolf remains have been found across a broad range of habitats including the plains, grasslands, and some forested mountain areas of North America and the arid savanna of South America. The sites range in elevation from sea level to 2,255 meters (7,400 ft). Dire wolf fossils have rarely been found north of 42°N latitude; there have been only five unconfirmed reports above this latitude. This range restriction is thought to be due to temperature, prey, or habitat limitations imposed by proximity to the Laurentide and Cordilleran ice sheets that existed at the time.

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    From the 1850s, the fossil remains of extinct large wolves were being found in the United States, and it was not immediately clear that these all belonged to one species. The first specimen of what would later become associated with Aenocyon dirus was found in mid-1854 in the bed of the Ohio River near Evansville, Indiana. The fossilized jawbone with cheek teeth was obtained by the geologist Joseph Granville Norwood from an Evansville collector, Francis A. Linck. The paleontologist Joseph Leidy determined that the specimen represented an extinct species of wolf and reported it under the name of Canis primaevus. Norwood's letters to Leidy are preserved along with the type specimen (the first of a species that has a written description) at the Academy of Natural Sciences of Philadelphia. In 1857, while exploring the Niobrara River valley in Nebraska, Leidy found the vertebrae of an extinct Canis species that he reported the following year under the name C. dirus. The name C. primaevus (Leidy 1854) was later renamed Canis indianensis (Leidy 1869) when Leidy found out that the name C. primaevus had previously been used by the British naturalist Brian Houghton Hodgson for the dhole.

    In 1876 the zoologist Joel Asaph Allen discovered the remains of Canis mississippiensis (Allen 1876) and associated these with C. dirus (Leidy 1858) and Canis indianensis (Leidy 1869). As so little was found of these three specimens, Allen thought it best to leave each specimen listed under its provisional name until more material could be found to reveal their relationship. In 1908 the paleontologist John Campbell Merriam began retrieving numerous fossilized bone fragments of a large wolf from the Rancho La Brea tar pits. By 1912 he had found a skeleton sufficiently complete to be able to formally recognize these and the previously found specimens under the name C. dirus (Leidy 1858). Because the rules of nomenclature stipulated that the name of a species should be the oldest name ever applied to it, Merriam therefore selected the name of Leidy's 1858 specimen, C. dirus. In 1915 the paleontologist Edward Troxell indicated his agreement with Merriam when he declared C. indianensis a synonym of C. dirus. In 1918, after studying these fossils, Merriam proposed consolidating their names under the separate genus Aenocyon (from ainos, 'terrible' and cyon, 'dog') to become Aenocyon dirus, but at that time not everyone agreed with this extinct wolf being placed in a new genus separate from the genus Canis. Canis ayersi (Sellards 1916) and Aenocyon dirus (Merriam 1918) were recognized as synonyms of C. dirus by the paleontologist Ernest Lundelius in 1972. All of the above taxa were declared synonyms of C. dirus in 1979, according to the paleontologist Ronald M. Nowak.

    In 1984 a study by Björn Kurtén recognized a geographic variation within the dire wolf populations and proposed two su…

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    The average dire wolf proportions were similar to those of two modern North American wolves: the Yukon wolf (Canis lupus pambasileus) and the Northwestern wolf (Canis lupus occidentalis). The largest northern wolves today have a shoulder height of up to 38 in (97 cm) and a body length of 69 in (180 cm). Some dire wolf specimens from Rancho La Brea are smaller than this, and some are larger.

    The dire wolf had smaller feet and a larger head when compared with a northern wolf of the same body size. The skull length could reach up to 310 mm (12 in) or longer, with a broader palate, frontal region, and zygomatic arches compared with the Yukon wolf. These dimensions make the skull very massive. Its sagittal crest was higher, with the inion showing a significant backward projection, and with the rear ends of the nasal bones extending relatively far back into the skull. A connected skeleton of a dire wolf from Rancho La Brea is difficult to find because the tar allows the bones to disassemble in many directions. Parts of a vertebral column have been assembled, and it was found to be similar to that of the modern wolf, with the same number of vertebrae.

    Geographic differences in dire wolves were not detected until 1984, when a study of skeletal remains showed differences in a few cranio-dental features and limb proportions between specimens from California and Mexico (A. d. guildayi) and those found from the east of the Continental Divide (A. d. dirus). A comparison of limb size shows that the rear limbs of A. d. guildayi were 8% shorter than the Yukon wolf due to a significantly shorter tibia and metatarsus, and that the front limbs were also shorter due to their slightly shorter lower bones. With its comparatively lighter and smaller limbs and massive head, A. d. guildayi was not as well adapted for running as timber wolves and coyotes. A. d. dirus possessed significantly longer limbs than A. d. guildayi. The forelimbs were 14% longer than A. d. guildayi due to 10% longer humeri, 15% longer radii, and 15% longer metacarpals. The rear limbs were 10% longer than A. d. guildayi due to 10% longer femora and tibiae, and 15% longer metatarsals. A. d. dirus is comparable to the Yukon wolf in limb length. The largest A. d. dirus femur was found in Carroll Cave, Missouri, and measured 278 mm (10.9 in).

    A. d. guildayi is estimated to have weighed on average 60 kg (132 lb), and A. d. dirus weighed on average 68 kg (150 lb) with some specimens being larger, but these could not have exceeded 110 kg (243 lb) due to skeletal limits. In comparison, the average weight of the Yukon wolf is 43 kg (95 lb) for males and 37 kg (82 lb) for females. Individual weights for Yukon wolves can vary from 21 kg (46 lb) to 55 kg (121 lb), with one Yukon wolf weighing 79.4 kg (175 lb). These figures show th…

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    Ecological factors such as habitat type, climate, prey specialization, and predatory competition have been shown to greatly influence gray wolf craniodental plasticity, which is an adaptation of the cranium and teeth due to the influences of the environment. Similarly, the dire wolf was a hypercarnivore, with a skull and dentition adapted for hunting large and struggling prey; the shape of its skull and snout changed across time, and changes in the size of its body have been correlated with climate fluctuations.
    The last glacial period, commonly referred to as the "Ice Age", spanned 125,000 –14,500 YBP and was the most recent glacial period within the current ice age, which occurred during the last years of the Pleistocene era. The Ice Age reached its peak during the Last Glacial Maximum, when ice sheets began advancing from 33,000 YBP and reached their maximum limits 26,500 YBP. Deglaciation commenced in the Northern Hemisphere approximately 19,000 YBP and in Antarctica approximately 14,500 YBP, which is consistent with evidence that glacial meltwater was the primary source for an abrupt rise in sea level 14,500 YBP. Access into northern North America was blocked by the Wisconsin glaciation. The fossil evidence from the Americas points to the extinction mainly of large animals, termed Pleistocene megafauna, near the end of the last glaciation.

    Coastal southern California from 60,000 YBP to the end of the Last Glacial Maximum was cooler and with a more balanced supply of moisture than today. During the Last Glacial Maximum, the mean annual temperature decreased from 11 °C (52 °F) down to 5 °C (41 °F) degrees, and annual precipitation had decreased from 100 cm (39 in) down to 45 cm (18 in). This region was unaffected by the climatic effects of the Wisconsin glaciation and is thought to have been an Ice Age refugium for animals and cold-sensitive plants. By 24,000 YBP, the abundance of oak and chaparral decreased, but pines increased, creating open parklands similar to today's coastal montane/juniper woodlands. After 14,000 YBP, the abundance of conifers decreased, and those of the modern coastal plant communities, including oak woodland, chaparral, and coastal sage scrub, increased. The Santa Monica Plain lies north of the city of Santa Monica and extends along the southern base of the Santa Monica Mountains, and 28,000–26,000 YBP it was dominated by coastal sage scrub, with cypress and pines at higher elevations. The Santa Monica Mountains supported a chaparral community on its slopes and isolated coast redwood and dogwood in its protected canyons, along with river communities that included willow, red cedar, and sycamore. These plant communities suggest a winter rainfall similar to that of modern coastal southern California, but the presence of coast redwood now found 600 kilometres (370 mi) to the nort…

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    Dire wolf remains have been found across a broad range of habitats including the plains, grasslands, and some forested mountain areas of North America, the arid savannah of South America, and possibly the steppes of eastern Asia. The sites range in elevation from sea level to 2,255 m (7,400 ft). The location of these fossil remains suggests that dire wolves lived predominantly in the open lowlands along with their prey the large herbivores. Dire wolf remains are not often found at high latitudes in North America, with the northernmost record in southern Canada.

    In the United States, dire wolf fossils have been reported in Arizona, California, Florida, Idaho, Indiana, Kansas, Kentucky, Missouri, Nebraska, New Mexico, Oregon, Pennsylvania, South Carolina, South Dakota, Texas, Utah, Virginia, West Virginia, Wyoming, and Nevada. The identity of fossils reported farther north than California is not confirmed. There have been five reports of unconfirmed dire wolf fossils north of 42°N latitude at Fossil Lake, Oregon (125,000–10,000 YBP), American Falls Reservoir, Idaho (125,000–75,000 YBP), Salamander Cave, South Dakota (250,000 YBP), and four closely grouped sites in northern Nebraska (250,000 YBP). This suggests a range restriction on dire wolves due to temperature, prey, or habitat. The major fossil-producing sites for A. d. dirus are located east of the Rocky Mountains and include Friesenhahn Cave, near San Antonio, Texas; Carroll Cave, near Richland, Missouri; and Reddick, Florida.

    Localities in Mexico where dire wolf remains have been collected include El Cedazo in Aguascalientes, Comondú Municipality in Baja California Sur, El Cedral in San Luis Potosí, El Tajo Quarry near Tequixquiac, state of Mexico, Valsequillo in Puebla, Lago de Chapala in Jalisco, Loltun Cave in Yucatán, Potrecito in Sinaloa, San Josecito Cave near Aramberri in Nuevo León and Térapa in Sonora. The specimens from Térapa were confirmed as A. d. guildayi. The finds at San Josecito Cave and El Cedazo have the greatest number of individuals from a single locality.

    In South America, dire wolves have been dated younger than 17,000 YBP and have been reported from six localities: Muaco in the western Falcón state of Venezuela, Talara Province in Peru, Monagas state in eastern Venezuela, the Tarija Department in Bolivia, Atacama Desert of Chile, and Ecuador. If the dire wolf originated in North America, the species likely dispersed into South America via the Andean corridor, a proposed pathway for temperate mammals to migrate from Central to South America because of the favorable cool, dry, and open habitats that characterized the region at times. This most likely happened during a glacial period because the pathway then consisted o…

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