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15.12.2015 16:55 - Encyclopedia Largest prehistoric animals Vol.1 Vertebrates part1 Mammals ch.1 Carnivores - Canine,sequel - Dogs and Wolves
Автор: valentint Категория: Забавление   
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Последна промяна: 03.07.2019 03:41


1. Borophagus 2. Cynarctus 3. Aelurodon 4. Osteoborus 5. Epicyon
Paleontologists agree that the late Eocene (about 40 to 35 million years ago) Hesperocyon was directly ancestral to all later canids—and thus to the genus Canis, which branched off from a subfamily of canids about six million years ago. This "western dog" was only about the size of a small fox, but its inner-ear structure was characteristic of later dogs, and there"s some evidence that it may have lived in communities, either high up in trees or in underground burrows. Hesperocyon is very well-represented in the fossil record; in fact, this was one of the most common mammals of prehistoric North America.
Another group of early canids was the borophagines, or "bone-crushing dogs," equipped with powerful jaws and teeth suitable for scavenging the carcasses of mammalian megafauna. The largest, most dangerous borophagines were the 100-pound Borophagus and the even bigger Epicyon; other genera included the earlier Tomarctus and Aelurodon, which were more reasonably sized. We can"t say for sure, but there"s some evidence that these bone-crushing dogs (which were also restricted to North America) hunted or scavenged in packs, like modern hyenas.

Here"s where things get a bit confusing. Shortly after the appearance of Hesperocyon 40 million years ago, Leptocyon arrived on the scene—not a brother, but more like a second cousin once removed. Leptocyon was the first true canine (that is, it belonged to the caninae subfamily of the canidae family), but a small and unobtrusive one, not much bigger than Hesperocyon itself. The immediate descendant of Leptocyon, Eucyon, had the good fortune to live at a time when both Eurasia and South America were accessible from North America—the first via the Bering land bridge, and the second thanks to the uncovering of central America. In North America, about six million years ago, populations of Eucyon evolved into the first members of the modern dog genus Canis, which spread to these other continents.
But the tale doesn"t end there. Although canines (including the first coyotes) continued to live in North America during the Pliocene epoch, the first plus-sized wolves evolved elsewhere, and "re-invaded" North America shortly before the ensuing Pleistocene (via that same Bering land bridge). The most famous of these canines was the Dire Wolf, Canis diris, which evolved from an "old world" wolf that colonized both North and South America (by the way, the Dire Wolf competed directly for prey with Smilodon, the "saber-toothed tiger.")
The end of the Pleistocene epoch witnessed the rise of human civilization around the world. As far as we can tell, the first domestication of the Gray Wolf occurred somewhere in Europe or Asia anywhere from 30,000 to 15,000 years ago. After 40 million years of evolution, the modern dog had finally made its debut!

The largest wild dog of all time was Epicyon haydeni, which stood 37 inches tall (0.9 meters) at the shoulder.
Epicyon ("more than a dog") is a large, extinct, canidgenus of the subfamilyBorophaginae ("bone-crushing dogs"), native to North America. Epicyon existed for about 15 million years from the Hemingfordian age of the Early Miocene to the Hemphillian of the Late Miocene.

Epicyon was about 5 feet long, and its weight has been estimated at 150 pounds. Epicyon had a massive head and powerful jaws, giving its skull a lion-like shape rather than that of a wolf.
Epicyon was one of the last of the Borophaginae and shared its North American habitat with other canids:
Borophagus23.3 to 3.6million years ago (mya)
Epicyon from 20.6 to 5.330 mya
Carpocyon from 20.4 to 3.9 mya
Paratomarctus from 16.3 to 5.3 mya
Aelurodon from 15.97 to 5.332 mya
Canis lepophagus from 10.3 to 1.8 mya.Canis lepophagus may be the ancestor to the wolf.


1. Eucyon 2. Leptocyon 3. Canis dirus 4. Tomarctus 5. Hesperocyon
How and when did wolves evolve? It"s a question science has yet to answer conclusively because, evolutionarily speaking, the development of the modern day wolf isn"t clear. What"s generally believed is that about 60 million years ago during what"s referred to as the Paleocene times wolves began their development as a specialized genus of cursorial, or hunt-by-chasing, carnivore. Approximately 8 million years later primitive carnivores known as Miacids appeared in the Lower Tertiary. The Tertiary period lasted about 65 million years and is often called "the age of the rise of mammals." Miacids ranged from gopher to dog size and had evolved from Cretaceous insectivores.
So, small rodent-like insectivores are included in the wolf"s ancestry. Much later, creodonts emerged. These animals are described as walking on five toes, having partially retractile claws, partially opposable thumbs on the forefeet and long thick tails. In his book Of Wolves And Men, Barry Holstun Lopez hypothesizes that creodonts "perhaps looked like long-legged otters, dwelt in forests, and may have slept in trees." Of course evolution continued and some creodonts eventually transformed. These transformations eventually moved out onto plains and prairies becoming wolves, bears, badgers, skunks and weasels. Those that remained behind in the forests became saber-toothed tigers, leopards and cheetahs.
By about 20 million years ago during what"s called the Miocene times, the two superfamilies of carnivores, canines and felines, were distinct. This is really when the more recognizable ancestors of the wolf emerged. Lopez describes them as having "specialized shearing teeth" with the bones of their lower legs having fused "as flexibility in the limbs...gave way for rigidity for strength in the chase." In Tomarctus, one ancestor of the wolf, the fifth toe on the hind leg became vestigial and the dew claw was born. Apparently Tomarctus lived much like the wild dogs and wolves of today. It"s described as a stealthy predator with short prick ears and a long tail that was used to maintain balance.
As Robert H. Busch outlines in his book The Wolf Almanac, research by Robert Wayne at the University of California suggests that a number of wolf-like canids diverged from a common ancestor about two to three million years ago. Early in the Pleistocene period, about a million years ago, the first Gray Wolf, Canis lupus, probably appeared in Eurasia.  It"s thought to have migrated to North America about 750,000 years ago. Evolving even earlier, the Dire Wolf, Canis dirus, was larger and heavier. It co-existed with the Gray Wolf in North America for approximately 400,000 years. Many believe that as the Dire Wolf"s main prey became extinct due to climactic change about 16,000 years ago the Dire Wolf itself eventually died out. Consequently, about 7,000 years ago, the Gray Wolf became the prime canine predator in North America. It was classified as Canis lupus by Swedish scientist Carl von Linne in 1758.
Why did Canis lupus survive while Canis dirus  did not? In his book The Great American Wolf, Bruce Hampton theorizes that since the Dire Wolf was "[built] more for power than speed and used to preying on relatively sedentary megaherbivores, dirus simply may have been unable to catch the smaller and faster ungulates that ascended as larger ones disappeared." In contrast, lupus was fast, with social behaviorism better equipped to meet the changing world. Hampton writes that perhaps the Dire Wolf was "less tolerant and more aggressive toward members of its own species."
Although the Dire Wolf appears to have hunted in large packs, these packs were probably less mobile and less far-ranging than those of the co-existing Gray Wolf. Ill-equipped to catch the faster prey that Gray Wolves lived upon, Hampton compares Dire Wolves to hyenas in that they may have resorted more and more to scavenging, "a survival strategy considered less versatile than that practiced by self-supporting carnivores." If in fact scavenging became the lifestyle of Dire Wolves, it proved insufficient to maintain their survival once megaherbivores, their original prey, completely died out.

The largest wolf is the dire wolf (Canis dirus) 1.5 m (4.9 ft) in length and weighed between 50 kg (110 lb) and 79 kg (174 lb).

The dire wolf (Canis dirus, "fearsome dog") is an extinctcarnivorousmammal of the genus Canis, roughly the size of the extant gray wolf, but with a heavier build. It evolved in North America and later moved into South America. Canis dirus is assigned to the Rancholabrean land mammal age of North America (240,000–10,000 years BP) and was among the many large carnivores and megaherbivores that became extinct in North and South America near the end of the Pleistoceneepoch. Salamander Cave in the Black Hills of South Dakota has produced the oldest known fossil of what is thought to be a dire wolf. The site preserves a fauna approximately 252,000 years old, based on uranium-series dating of a horse fossil found there. Its ending is associated with the Quaternary extinction event.
The dire wolf averaged about 1.5 m (5 ft) in length and weighed between 50 and 79 kg (110 and 174 lb), which made it the largest species in the genus Canis. Limb elements are rarely found outside the La Brea Tar Pits, which makes it hard to compare the size of average individuals between populations. The dire wolf is estimated to have been 8% shorter at the shoulder than the modern Northwestern wolf, and of equal height to the typical gray wolf, but more heavily built. With the exception of the canine teeth in some populations, male and female body and teeth sizes evidence no major sexual dimorphism, similar to most canines. In some populations, males’ canine teeth were considerably larger, suggesting male competition for breeding access. In other populations, lack of dimorphism in the canine teeth suggests little competition.
Despite superficial similarities to the gray wolf, the two species differed significantly. Today’s largest gray wolves would have been of similar size to an average dire wolf; the largest dire wolves would have been considerably larger than any modern gray wolf. The dire wolf is calculated to have weighed 25% more than living gray wolves.
Many of these characteristics were needed to fight off and prey on larger megafauna. The legs of the dire wolf were proportionally shorter and sturdier than those of the gray wolf, and its brain case was smaller than that of a similarly sized gray wolf.
The dire wolf"s teeth were similar to the gray wolf"s, only slightly larger, pointing to a hypercarnivorous to mesocarnivorous activity. The dietary characteristics were primarily carnivorous, as well as partially omnivorous.
Many paleontologists have proposed that the dire wolf may have used its relatively large teeth to crush bone, an idea supported by the frequency of large amounts of wear on the crowns of their fossilized teeth. The upper carnassial had much larger blade than that of the gray wolf, indicating greater slicing ability. It had a longer temporal fossa and broader zygomatic arches, indicating the presence of a large temporalis muscle capable of generating slightly more force than a gray wolf"s. However, other scientists have noted the dorsoventral and labiolingual force profiles are indistinguishable from those of other canids, such as coyotes and African wild dogs, indicating a similar diet.
Dire wolf teeth lacked the craniodental adaptations of habitual bonecrushers such as hyenas and borophagines. The dire wolf"s robust skull and susceptibility to tooth breakage suggests that it hunted for large prey or scavenged the carcasses of large prey. The dorsoventrally weak symphyseal region indicates it killed in a manner similar to its modern relatives, by delivering a series of shallow bites, strongly indicating pack hunting behaviour. However, the incidence of broken postcarnassial molars is much higher than in fossil gray wolves, indicating the species was probably less adapted to bone crushing than the gray wolf.
Dire wolves are part of the same carnivorous guild as the smaller gray wolves and coyotes.Dire wolves" overpowering bite, 129% of the force of the modern gray wolf, could hold and subdue their prey. As inferred from their large bodies and carnivorous teeth, they often took on large prey or megafauna, made possible by traveling in packs. Dire wolves were not specialized hunters—they fed on whatever megafauna was abundant.
Compared to modern species, a remarkable number of dire wolf specimens from the La Brea pits showed evidence of having broken their teeth in life. Specimens in the older part of the pit exhibited more tooth wear than those in the younger pit, which could be a result of either the older part of the pit containing more senior dire wolf specimens or a diet that included harder foods such as carcasses and bones.Another explanation for increased tooth wear and breakage could be intense competition among carnivores forcing C. dirus to eat as much of the carcass as quickly as possible.
The dire wolf has an extremely extensive fossil range. It is known to be located as far north as Canada and south into southern Bolivia. Ten localities in Mexico are known to contain Canis dirus: Cedazo, Aguascalientes; Comondu, Baja California; El Cedral, San Luis Potosн; El Tajo Quarry, Tequixquiac, and Valsequillo, all in Distrito Federal, Mexico; Lago de Chapala, Jalisco; Loltun Cave, Yucatбn; Potrecito, Sinaloa; and San Josecito Cave, Nuevo Leуn. The majority of these localities occur in the central and southeast-central portions of Mexico with few occurrences in the north or west. Of the central localities, San Josecito Cave and Cedazo have the greatest number of individuals of Canis dirus collected from a single locality; other localities in Mexico are known by only a few specimens
Due to the large amount of remains at the Rancho La Brea tar pits it is possible dire wolves used animals trapped here as a food source and some end up being trapped and preserved themselves. It can also be concluded that they were social predators as they are most successful in defending carcasses.
The habitat of C. dirus varied considerably. In North America, it ranged from plains and grasslands to forested mountain areas. In South America, it occupied areas of arid savannah. The dire wolf lived in several habitats, tropical marsh with thorn-scrub to deciduous forest including some component of nearby grassland, and from sea level up to 2255 m (7400 feet). It was widespread, and its remains have been found in 136 places, from Alberta, Canada, to Tarija, Bolivia. Although the dire wolf was recorded as far north as Alberta, its remains have never been found at higher latitudes. The dire wolf may have migrated to northern Canada during interglacial periods, but its remains would likely have been obliterated by later glacial activity. However, the last named species of dire wolf was said to be found in North America stemming from Florida. The largest known dire wolves specimens have also said to have been recovered from this area of North America. So even though the dire wolf has migrated it may have found to be seasonal.
The dire wolf is well known for its unusually high representation in La Brea Tar Pits in California. Over 200,000 fragments representing more than 4,000 individual dire wolves have been recovered from the tar pits, more than any other mammal species. This large number suggests the dire wolf, like modern wolves and dogs, hunted in packs. The abundance of remains of the gray wolf (C. lupus, also known as C. furlongi) in the tar pits is about 1% that of the dire wolf.
Whether the dire wolf originated in North America versus South America is the subject of controversy. Most paleontologists lean toward a North American origin for three reasons: first, more potential progenitors are present in the middle Pleistocene of North America; second, distribution of C. dirus is much better represented in North America, with 136 sites versus only three localities in South America; and last, C. dirus appears earlier in the fossil record in North America than South America. A North American origin implies that C. dirus migrated into South America from North and Central America. The dire wolves most likely migrated to 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, however, as the pathway then consisted of open, arid regions and savanna whereas during inter-glacial periods, it would have been characterized by tropical rain forest habitat.
The fossil record suggests that the genus Canis diverged from the small, foxlike Leptocyon in North America sometime in the Late Miocene epoch 9 to 10 million years (Ma) ago, along with two other genera, Urocyon, and Vulpes. Canids soon spread to Asia and Europe (8 Ma BP) and became the ancestors of modern wolves, jackals, foxes, and the raccoon dog.
By 3–5 Ma BP, canids had spread to Africa (Early Pliocene) and South America (Late Pliocene). Their invasion of South America as part of the Great American Interchange was enabled by the formation of the Isthmus of Panama 3 Ma ago.
Over the next nine million years, extensive development and diversification of the North American wolves took place by the Middle Pleistocene. Canis armbrusteri appeared, possibly from C. chihliensis in Asia. There is good evidence that the dire wolf evolved from C. armbrusteri, with the two taxa sharing in the open plains and grasslands of what is now the central United States.
C. dirus eventually displaced C. armbrusteri, with the latter"s final range shrinking to what is now the southeastern U.S., more specifically Florida. While this occurred, C. dirus expanded its range to include that of C. armbrusteri and moved into Central America and South America, appearing in the Late Pleistocene fossil record in northwestern South America.
Two subspecies of the dire wolf are known to have inhabited what is now the United States. C. dirus guildayi was smaller and ranged west of the Rocky Mountains. C. dirus dirus was larger and ranged east of the Rockies.
In 2010, a genetic study of extinct South American canids indicated that the South American Canis nehringi and Canis dirus were the same species, with the Gray wolf Canis lupus being in a sister clade.
Although it was closely related to the gray wolf and other sister species, C. dirus is not the direct ancestor of any modern species. Unlike the gray wolf, which is of Eurasian origin, the dire wolf evolved on the North American continent, along with the coyote. The dire wolf co-existed with the gray wolf in North America for about 100,000 years.
The dire wolf was one of the abundant Pleistocene megafauna—a wide variety of large mammals that lived during the Pleistocene. Approximately 10,000 years ago the dire wolf became extinct along with most other North American megafauna.
During the Late Pleistocene (300,000 years ago) the gray wolf (C. lupus) crossed into North America on the Bering Strait land bridge and competed with the dire wolf. Overlapping fossil findings of the extinct saber tooth cat, the Smilodon, shows that the dire wolf had these as competition in North America. Both species were social animals that hunt in packs and preyed on the same animals. Starting about 16,000 years ago, coinciding with the end of the last glacial period and the arrival of humans in North America, most of the large mammals upon which the dire wolf depended for prey began to die out, possibly as a result of climate and/or human-induced changes.
Slower than the other wolf species on the continent at the time, like the gray wolf and red wolf, the dire wolf could not hunt the remaining swifter prey, and was forced to subsist on scavenging. By approximately 10,000 years ago, the large mammals and the dire wolf were extinct. Their demise, along with that of other large Pleistocene carnivores, was found to be related to the extinction of megafaunal prey.
More information is required in order to understand the extinction of C. dirus, including information on its biogeographical range, population size, competition, interactions with predators and prey, and its physical environment. Further discoveries about the dire wolf"s interactions with its competition and prey around the time of the extinction events would also be integral in understanding why C. dirus went extinct.




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