Encyclopedia Largest prehistoric animals Vol.1 Vertebrates part1 
Mammals ch.3 Carnivores - Canine,from the huge bear-dogs to dire wolf
The largest bear-dog was a species of Pseudocyon weighing around 773 kg, representing a very large individual

Pseudocyon, also known as Amphicyonopsis, is an extinct genus of terrestrialcarnivores belonging to the suborderCaniformia, family of Amphicyonidae ("bear dog") and which inhabited Euroasia and North America from the Miocene epoch to the Late Miocene subepoch living 23.3—7.2 Mya. Pseudocyon existed for approximately 5.3 million years.
With an upper weight estimate exceeding seven hundred and seventy kilograms, Pseudocyon was one of the largest of the bear dogs, quite probably even bigger than the largest known specimen of Amphicyon
 which was estimated to come from a six hundred kilogram individual. 
Additionally Pseudocyon also had a similar geographic distribution to 
Amphicyon although Pseudocyon does seem to have lived for slightly 
longer during the Miocene.
Pseudocyon was assigned to Amphicyoninae by Hunt in 1988 
and to Amphicyonidae by Lartet (1851), Carroll (1988) and Pickford et 
al. in 2000. It is synonymous with Amphicyonopsis.
Specimens were located in Belomechetskaia Russian Federation, Santa Cruz, New Mexico, Pontigne and Malartic, a la ferme Larrieu, France, and Nebraska. The largest fossil find was of a mandible
 (F:AM 49247) founded in New Mexico. The mass estimate derived from the 
mandible was about 773 kg, representing a very large individual of 
Pseudocyon.
Very close contender is Amphicyon giganteus
 Size: Up to 2.5 meters long. 1,4 meters tall at the shoulder.Weight estimated at up to  750 kg.
Amphicyon giganteus is an extinct species of Amphicyon, a large 
carnivorous bone cruhing mammal known as a Bear-Dog of the family 
Amphicyonidae, subfamily Amphicyoninae, from the Miocene endemic to 
Europe and Africa, living from 16.8—7.2 Ma existing approximately 9.7 
million years An account of an ancient Rhinocerotidae (Iberotherium 
rexmanueli zbyszewskii) which may have been killed by a single 
individual or by a pack of Amphicyon giganteus in now what is Portugal, 
was written noted paleontologists Ginsberg et al. Amphicyon castellanus 
also shared its time period and habitat. Amphicyon giganteus was named 
by Kaup (1884). Amphicyon giganteus was the typical Bear-Dog Amphicyonid
 with morphology to both bears and dogs. Looking more like a bear than a
 dog, it had a large heavy tail, thick neck, robust limbs and teeth like
 a wolf. It was probably an omnivore with a lifestyle comparable to that
 of the brown bear
Size: Up to 2.5 meters long. 1,4 meters tall at the shoulder.Weight estimated at up to  750 kg.
Amphicyon giganteus is an extinct species of Amphicyon, a large 
carnivorous bone cruhing mammal known as a Bear-Dog of the family 
Amphicyonidae, subfamily Amphicyoninae, from the Miocene endemic to 
Europe and Africa, living from 16.8—7.2 Ma existing approximately 9.7 
million years An account of an ancient Rhinocerotidae (Iberotherium 
rexmanueli zbyszewskii) which may have been killed by a single 
individual or by a pack of Amphicyon giganteus in now what is Portugal, 
was written noted paleontologists Ginsberg et al. Amphicyon castellanus 
also shared its time period and habitat. Amphicyon giganteus was named 
by Kaup (1884). Amphicyon giganteus was the typical Bear-Dog Amphicyonid
 with morphology to both bears and dogs. Looking more like a bear than a
 dog, it had a large heavy tail, thick neck, robust limbs and teeth like
 a wolf. It was probably an omnivore with a lifestyle comparable to that
 of the brown bear
Amphicyon ingens is other very large bear-dog

Dimensions:length - 2,5 m, height - 120 сm, weight - 200-550 kg.
Amphicyon ("ambiguous dog") is an extinct genus of large carnivorous 
bone-crushing mammals, known as bear-dogs, of the family Amphicyonidae, 
subfamily Amphicyoninae, from the Aquitanian Epoch until the Tortonian. 
They ranged over North America, Europe, Asia, and Africa from 20.6—9 Ma 
ago, existing approximately 11.6 million years. Amphicyon was the 
typical bear-dog amphicyonid with morphology similar to both bears and 
dogs. With its robust build and maximum length of 2.5 m, the largest 
species looked more like a bear than a dog. It had a large heavy tail, 
thick neck, robust limbs and teeth like a wolf. It was probably an 
omnivore with a lifestyle comparable to that of the brown bear.
Amphicyon ingens lived from 20.06–13.6 Ma, approximately 6.46 
million years. The species was originally described by W. Matthew in 
1924 from specimens found in the Olcott Formation, Sioux County, 
Nebraska. Specimens attributed to this species have since been found in 
California, Colorado, and New Mexico. The largest known specimen to 
weigh 600 kg, making it the largest amphicyonid and one of the largest 
known carnivorous land mammals.
The largest canid 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.
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.
 Other than size, there is no significant special adaptations in the 
mandibular corpus of the dire wolf to set it apart from the gray wolf.
Between C. lupus and C. dirus, C. lupus had a significantly 
greater mechanical advantage of the temporalis muscle over C. dirus at 
the lower carnassial and P4 (MATM1, MATP4).
The slicing teeth (P4, the carnassial) on the upper jaw of C. 
dirus are larger than those of the gray wolf, but those on the lower jaw
 are similar. The temporalis of the dire wolf could generate more force 
than seen in modern gray wolves, suggesting stronger killing bites.
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.
The type specimen of the dire wolf was found in Evansville, Indiana, in the summer of 1854, when the Ohio River was quite low. The specimen, a fossilized jawbone, was obtained by Joseph Granville Norwood
 from an Evansville collector named Francis A. Linck.Norwood, who at 
that time was the first state geologist of Illinois, sent the specimen 
to Joseph Leidy at the Academy of Natural Sciences in Philadelphia.Leidy
 determined the specimen represented an extinct species of wolf and 
published a note to that effect in November 1854. In a publication dated
 1858, Leidy assigned the name Canis dirus.
Norwood"s letters to Leidy are preserved along with the type 
specimen at the Academy of Natural Sciences, although one of the letters
 indicates the specimen was to be returned to Linck"s family, as Linck 
died in August 1854.
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.
 
 
 
Encyclopedia Largest prehistoric animals Vol.1 Vertebrates part1 Mammals ch.3 Carnivores-Giant prehistoric bears,sequel
Cave bear- Ursus spelaeus

Ursus spelaeus, better known as the cave bear is by far one of the most 
common Pleistocene mammals in the fossil record, thanks mainly due to 
its behaviour of frequenting caves. In fact the fossils of Ursus 
spelaeus are so numerous that in World War I the German army used them 
as a source of phosphates.
The huge numbers of cave bear fossils in caves that have been documented
 to represent thousands of individuals to a single cave strongly 
suggests that this bear regularly spent time in caves, perhaps returning
 to rest after foraging. Bears that we know today by contrast only 
frequent caves during the hibernation period and sleep outdoors during 
the warmer months. The immense number of Ursus spelaeus fossils in 
certain caves has also led to the theory that these bears may have 
actually lived in social groups, although in depth study of the layers 
and dating has suggested that the remains are of single individual bears
 building up over the course of the Pleistocene period. As older remains
 became buried by sediment and fossilised they would have been of no 
nutritional value to anything and would have been left alone to 
accumulate, giving the impression of several bears living together 
rather than just one or two.
Cave bears have long been thought to be herbivorous animals, meaning 
that they only ate plants. Key evidence for this comes from the lack of 
premolar teeth that are usually absent in herbivores resulting in a gap 
between the forwards canines and rear molars called a diastema. Isotope 
analysis has also yielded low amounts of carbon-13 and nitrogen-15 which
 are usually found in high concentrations in carnivores. Counter to the 
strictly herbivorous diet theory however is the presence of some fossils
 that do have elevated levels of nitrogen-15, as, well as tooth wear 
associated with the gnawing of bones. Additionally some cave bear 
fossils also have tooth marks on them that seem to have been caused by 
others of their species. Not only does this suggest that cave bears 
occasionally ate meat, they may have scavenged the dead bodies of other 
cave bears that had died in caves. This is why Ursus spelaeus is today 
considered by most to have been an opportunistic omnivore that relied 
mostly upon eating plants, but would also supplement its diet with 
occasional meat when it was able to.
As one might expect cave bears were most common in areas that had large 
amounts of caves, particularly limestone that would have had caves 
carved out by water over several eons. As such cave bears lived in 
ecosystems that were between lowland plains and high level mountains 
that would have had a greater variety of trees and vegetation growing in
 them. Aside from offering a greater variety of food, a habitat 
preference such as this suggests that Ursus spelaeus did not compete 
with other European megafauna such as the woolly rhino and the woolly 
mammoth since they inhabited more open areas. Cave bears could also 
range across most of Europe, settling where they encountered suitable 
habitats, because the lower sea levels meant that marine boundaries such
 as the English Channel and North Sea did not exist.
A greater determining factor in the range of cave bears was the amount 
of glacial cover, something that would have expanded and receded over 
varying periods. Ursus spelaeus individuals from both milder and harsher
 times are known, with the individuals present in harsher times growing 
slightly larger than the bears that lived in slightly warmer times. This
 is a simple strategy where growing bigger creates a greater level of 
outer insulation while the total surface area of the body still 
increases but at proportionately less amount than the total body mass. 
This has been repeated and documented many times for different mammals 
that live in colder climates that all grow proportionately bigger that 
their closest related genera, perhaps the most relevant example here 
being the polar bear (Ursus maritimus) which does well in arctic 
conditions, but has been seen to overheat when kept in zoos in warmer 
climates.
Returning to the large collections of remains mentioned above, it’s 
clear that large numbers of Ursus spelaeus died in caves, but this needs
 further investigation. Because the remains of cave bears accumulated 
over tens of thousands of years it is easy to immediately discount the 
possibility of cave ins, as such an event would have only trapped one 
bear and then prevented others from entering. One widely accepted theory
 is that cave bears that did not eat enough food during the summer 
months to build up their fat reserves ended up starving to death as they
 slumbered. Even today animals that rely upon hibernation to last out 
the colder winter are vulnerable in times where plants do not grow or 
produce as much sustenance as they usually do, and back in the 
Pleistocene when ice sheets were toing and froing across Europe food 
would have been even more unpredictable from year to year.
More in depth study of some cave bear remains has revealed that bone 
disorders such as rickets, periostitus and osteomyelitis were quite 
common including other ailments such as the presence of tumours. 
Although not necessarily fatal to the individuals these bones belonged 
to, they would have impeded their ability to forage, slowly bringing the
 animal down to the point where it could no longer support itself. In 
these weakened states its thought that cave bears may have even fallen 
as prey to cave hyenas as well as even the European cave lion, that 
otherwise may have given a healthy bear in the prime of its life a wide 
birth.
Ultimately as a species, cave bears seem to have succumbed to the 
effects of habitat loss, as by only living in caves these bears would 
have only ever had a set amount of areas available to them. However as 
the Pleistocene was reaching its final stages, Neanderthals were 
beginning to become more common, and these primitive people also used 
caves for shelter. Inevitably one species would have to give way, and it
 was the cave bear that lost out to the greater numbers and intelligence
 of Neanderthals.
However despite the fact that they seem to have taken over, Neanderthals
 also seem to have held cave bears in very high regard. There are 
several burial sites in Europe where the remains of several bears have 
been assembled in pits and then covered with stone slabs. Perhaps the 
most famous site is Drachenloch, Switzerland where seven cave bear 
skulls are arranged to face the front of the cave, while six more are 
placed in recessed notches in the cave wall further in. Further remains 
were found bundled together, along with a skull of a three year old 
juvenile bear that had had its cheek pierced by the leg bone of another 
juvenile bear. Although some researchers claim that these are natural 
occurrences, there are a great many other who believe that remains like 
these are those of an ancient bear cult. How and why bears would be 
worshipped is uncertain, but it could be for anything from a totem 
animal, to a guardian of the caves against intruders, to maybe even a 
ward against other cave bears wandering in to Neanderthal settlements.
Ursus deningeri

Ursus deningeri (Deninger"s bear) is an extinct species of mammal 
of the family Ursidae (bears), endemic to Eurasia during the Pleistocene
 for approximately 1.7 million years, from ~1.8 Mya to 100,000 years 
ago. This large contemporary with the Cave Bear is reached 
sizes from 2.5 meters to length , 1.4 meters in height at the withers 
and weigh up to 600 kilograms.The range of this bear has 
been found to encompass both Europe and Asia, demonstrating the ability 
of the species to adapt to many Pleistocene environments. U. deningeri 
is a descendant of U. savini and an ancestor of U. spelaeus.
Ursus deningeri has a combination of primitive and derived 
characters that distinguishes it from all other Pleistocene bears. Its 
mandible is slender like that of living brown bears and Ursus etruscus. 
It also has derived characters of cave bears (Ursus spelaeus) and is 
considered to be the descendant of Ursus savini and very close to the 
common ancestor of brown bears.
Ursus ingressus

This large contemporary with the Cave Bear is reached sizes from 
2.3 meters to length , 1.3 meters in height at the withers and weigh up 
to 350 kilograms.
Whereas U. spelaeus inhabited mainly low and medium elevation 
areas, U. ingressus has mostly been found in medium and high elevated 
regions . Recent isotopic analyses showed also some dietary 
differentiation between these cave bear haplogroups. Ursus ingressus was
 likely better adapted to continental environments and, thus, might have
 outperformed U. spelaeus during cold and arid climate conditions.
The vast majority of fossil remains in Late 
Pleistocene deposits from Niedźwiedzia Cave in Kletno, Sudetes, Poland, 
belong to the cave bear. Phylogenetic analyses based on a fragment of 
the mitochondrial D-loop region extracted from two cave bear samples 
unambiguously showed their close relationship with the Ursus ingressus 
haplogroup. This taxonomic affiliation of the cave bear remains from 
Niedźwiedzia Cave was further confirmed by biometrical analyses of molar
 teeth and skulls. Our results represent the first record of U. 
ingressus north of the Carpathian Arch, while radiocarbon dating (> 
49,000 yr BP) of the samples indicates that they represent some of the 
oldest specimens of this cave bear taxon known so far. Multi-method 
phylogenetic analyses including numerous publicly available cave bear 
sequences allowed analysing the relationships among these samples in 
details, including the significance of particular clades, and discussing
 some aspects of cave bear phylogeography. The sequences of U. ingressus
 from Poland are most closely related to specimens from the Ural 
Mountains and next to Slovenia, which may indicate migrations between 
Central and Eastern European populations. The internal placement of 
Ural.   
Florida cave bear - Tremarctos floridanus

                                                  
Tremarctos floridanus occasionally called the Florida spectacled bear, 
Florida cave bear, or rarely Florida short-faced bear, is an extinct species of bear in the familyUrsidae,subfamily Tremarctinae. T. floridanus was endemic to North America from the Pliocene to Holocene epoch (4.9 million — 11,000 years ago), existing for approximately 4.9 million years.Tremarctos floridanus.
This smaller contemporary giantess bears Arktodus and Arctotherium weighed up to 450-500 kg.
T. floridanus was widely distributed south of the continental ice sheet, along the Gulf Coast across through Florida and north to Tennessee, and across the southern United States to California.
Arctodus (3 million — 11,000 years ago) was a 
contemporary and shared its habitat with T. floridanus. The closest 
living relative of the Florida cave bear is the spectacled bear of South America; they are classified together with the huge short-faced bears in the subfamilyTremarctinae. They became extinct at the end of the last ice age, 10,000 years ago (possibly as late as 8,000 years ago at Devil"s Den in Florida), due to some combination of climate change and hunting by newly arrived Paleo-Indians.
 
 
 
 
Encyclopedia largest prehistoric animals Vol.1 Vertebrates part1 Mammals ch.3 Carnivores-Bears,fearsome giants atop the food chain on land
Attempted extended overview of the largest ever lived 
animals.Codex consists of collected and processed by me but written by 
other authors articles. Full overview of vertebrates including the 
latest paleontological finds. Because of the volume "ve split up. I 
begin with predatory mammals because these are my favorites.Individual 
parts are numbered according to their place in the full collection 
Proceedings in English in order to reach the maximum number of readers.And to preserve the style of the authors.
 
Опит за разширен обзор на най-големите,живели някога животни.Сборника
 се състои от събрани и обработени от мен,но написани от други автори 
статии.Пълен обзор на гръбначните животни,с включване най-новите 
палеонтологични находки.Заради обема съм го разделил на части.Започвам с
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Bears
 Arctotherium
 
Arctotherium

The largest terrestrial carnivoran and the largest bear as well as the largest mammalian land-predator of all time was Arctotherium angustidens of the genus Arctotherium or the South American short-faced bears. A humerus of A. angustidens from Buenos Aires
 indicate that the big males of this species would have weighed 1,588- 
1,749 kg and standing at least 11 feet (3.4 meters) tall on the 
hind-limbs.Arctotherium angustidens was a South-American Short Faced Bear from the genus Arctotherium. It was the largest Carnivoran that ever lived, in most regards; although male southern elephant seals can be heavier, they are semi-aquatic and covered in a layer of blubber.

A. angustidens was the largest Carnivoran that ever lived 
except that for southern and northern elephant seals. The northern 
elephant seal was 3,700 kg (8,200 lb) while the southern elephant seal 
was 5,000 kg (11,000 lb). It weighted about 983–2,042 kg 
(2,167–4,502 lb) but the authors said it was more likely 1,588 kg 
(3,501 lb) Arctotherium is an extinct genus of South American 
short-faced bears within Ursidae of the Pleistocene. Their ancestors migrated from North America to South America during the Great American Interchange, following the formation of the Isthmus of Panama during the late Pliocene. The oldest remains are from the Ensenadan epoch within the early-middle Pleistocene 1.2 Mya. Their closest relatives were the North American short-faced bears of genus Arctodus (A. pristinus and A. simus). The closest living relative would be the spectacled bear (Tremarctos ornatus)
Arctotherium was named by Hermann Burmeister in 1879. It was assigned to Tremarctinae by Krause et al. 2008. A specimen of A. angustidens from Buenos Aires
 shows an individual estimated, using the humerus, to weigh between 983 
and 2,042 kg (2,167 and 4,502 lb), though the authors consider the upper
 limit as improbable and say that 1,588 kg (3,501 lb) is more likely. It
 is still possibly the largest bear ever found and contender for the 
largest carnivorous land mammal known to science. In contrast to their 
North American relatives, South American short-faced bears showed a 
trend of declining size and carnivory over time. This has been attributed to increased competition from other, later-arriving or evolving carnivorans, like jaguars, lions or Smilodon populator,
 following the early dispersal of short-faced bears to South America. 
(The North American carnivorans that invaded South America, including 
short-faced bears and Smilodon, quickly dominated the predatory niches formerly occupied by South America"s native metatheriansparassodont and avian phorusrhacid carnivores.)
North American short-faced bear (Arctodus simus)
 
The short-faced bear (Arctodus spp.) is an extinct bear that inhabited North America during the Pleistocene epoch from about 1.8 Mya until 11,000 years ago. It was the most common early North American bear and was most abundant in California.
 There are two recognized species: Arctodus pristinus and Arctodus 
simus, with the latter considered one of the largest known terrestrial 
mammalian carnivores.
The name short-faced bear derives from the shape of their skulls, which 
appear to have a proportionally short snout compared to other bears; 
this characteristic is also shared by its extant relatives, the spectacled bear and the grizzly bear.
 However, this apparent shortness is an optical illusion caused by their
 deep snouts and short nasal regions. The scientific name of the genus, 
Arctodus, derives from the Greek language and means "bear tooth".
The short-faced bear belongs to a group of bears known as the Tremarctinae, which appeared in the Americas during the earliest parts of the late Miocene epoch in the form of Plionarctos, a genus considered ancestral to Arctodus, Arctotherium and the modern spectacled bear
 (Tremarctos ornatus). Although the early history of Arctodus is poorly 
known, it evidently became widespread in North America by the Kansan age about 800,000 years ago.
Arctodus simus first appeared during the middle Pleistocene in North America, about 800,000 years ago, ranging from Alaska to Mississippi,
 and it became extinct about 11,600 years ago. Its fossils were first 
found in the Potter Creek Cave, Shasta County, California. It might have
 been the largest carnivorous land mammal that ever lived in North 
America. Only one Giant Short-faced Bear skeleton has been found in 
Indiana, unearthed south of Rochester on west of Nyona Lake on Chet 
Williams" farm. It has become well known in scientific circles because 
it was the biggest most-nearly complete skeleton of a giant short-faced 
bear found in America. The original bones are in the Field Museum, 
Chicago. The new Indiana State Museum, Indianapolis, and the Yukon 
Beringia Interpretive Centre, Whitehorse, Yukon Territory, Canada, have 
casts made of the bones. In a recent study, the mass of six specimens 
was estimated, one-third of them weighed about 900 kg (1 short ton), the
 largest being UVP 015 at 957 kg (2,110 lb), suggesting specimens that 
big were probably more common than previously thought. It stood 8–10 
feet (2.4–3.0 m) tall on hind legs while a large specimen would have 
been 11–12 feet (3.4–3.7 m) tall with a 14 foot (4.3 m) vertical arm 
reach, and 5–6 feet (1.5–1.8 m) high at the shoulder when walking on all
 fours, it was tall enough to look a man in the eye. At Riverbluff Cave,
 Missouri, a series of claw marks up to 15 feet (4.57 m) high have been 
found along the cave wall indicating Short-faced bears over 12 feet 
(3.65 m) tall.
Arctodus pristinus inhabited more southerly areas, ranging from northern Texas to New Jersey in the east, Aguascalientes, Mexico to the southwest, and with large concentrations in Florida, the oldest from the Santa Fe River 1 site of Gilchrist County, Florida paleontological sites.
Researchers disagree on the diet of Arctodus. Analysis of their bones showed high concentrations of nitrogen-15,
 a stable nitrogen isotope accumulated by meat-eaters, with no evidence 
of ingestion of vegetation. Based on this evidence, A. simus was highly carnivorous and as an adult would have required 16 kg (35.3 lb) of flesh per day to survive.
One theory of its predatory habits envisages A. simus as a brutish
 predator that overwhelmed the large mammals of the Pleistocene with its
 great physical strength. However, despite being very large, its limbs 
were too gracile for such an attack strategy. Alternatively, long legs 
and speed (50–70 km/h (30–40 mph)) may have allowed it to run down 
Pleistocene herbivores, such as wild horses and saiga antelopes, in a cheetah-like
 fashion. However, in this scenario, the bear’s sheer physical mass 
would be a handicap. Arctodus skeletons do not articulate in a way that 
would have allowed for quick turns, an ability required of any predator 
that survives by killing agile prey. Paul Matheus, paleontologist at the
 University of Alaska Fairbanks, determined that Arctodus" moved in a pacing motion like a camel,
 horse, and modern bears, making it built more for endurance than for 
great speed. A. simus, according to these arguments, was ill-equipped to
 be an active predator, leading some to conclude that it was a kleptoparasite, using its enormous size to intimidate smaller predators, such as dire wolves, Smilodon, and American lions, from their kills.
Some authors also suggest that the giant short-faced bear and the cave bear were omnivores, like most modern bears, and the former may have eaten plants depending on availability.
Ursus maritimus tyrannus

Ursus maritimus tyrannus was a very large fossil subspecies of 
the polar bear that descended from an Arctic population of brown bears. 
Its name in Latin means Tyrant Sea Bear.
Ursus maritimus tyrannus was the first polar bear and evolved sometime 
in the mid-Pleistocene. While the oldest fossil is 100,000 years old, 
they are thought to have evolved between 100,000 and 250,000 years ago 
from a population of brown bears likely isolated by glaciation. That 
population is believed to have diminished in numbers quickly into a much
 smaller population selected by species individual variation who adapted
 better to the changed environment. Over time with intense selective 
pressures on a small population they evolved the characteristics of the 
first polar bears.
Initially the isolated brown bears were no different than the variations
 of brown bears of that time period. Because litters of cubs can show 
significant species variations in hair color and hair thickness, this 
gave certain individuals a survival advantage passed on each generation.
 Eventually skull changes and even changes in dentition occurred leading
 to the smooth and rather quick evolution of U. maritimus tyrannus.
U. maritimus tyrannus was considerably larger then its modern relative. 
If everything is scaled out correctly from its remains, it would had 
been 183 cm (6 ft) at the shoulders, 3,6 m (12 ft) long and would have 
weighted an average of 1.2 tons, making it the largest bear "and one of 
the largest mammalian carnivores to ever walk on land". Its tremendous 
size makes it even bigger than the other "largest" mammalian carnivores 
that ever lived, including Andrewsarchus, Agriotherium, and Arctodus 
simus. It"s speculated that this gigantic bear would, due to its 
formidable size and strength, have preyed on mammoths which also lived 
during the time
Agriotherium

Agriotherium size is  approximately 2.7 meters long and up to around 650 kilograms.
 
Another thing to consider is that if Agriotherium was a 
scavenger then it was likely getting to carcasses after all of the 
choice pieces of meat had been consumed with perhaps only bones being 
left. This would probably not be enough to thwart Agriotherium from a 
meal however since the short snout, strong jaw closing muscles and 
robust construction of the skull and jaws were all the things that 
Agriotherium needed to develop massive bite force. Computer modelling in
 a 2012 study (see links below) confirmed that Agriotherium had one of 
the largest bite forces known amongst the members of the Carnivora (A 
group of mammals that includes dogs, bears, cats, pinnipeds etc which 
are specially adapted to exist by eating meat). By being able to crack 
open bones, Agriotherium could access and eat the bone marrow within, 
and for those not familiar, bone marrow is one of the most nutritious 
parts of an animal, and can last for several years after an animals 
death when encased inside of the bones. One of the better known bears in
 the worlds fossil record, the Agriotherium genus is also easily one of 
the largest currently known. With this large size it would be tempting 
to portray Agriotherium as a savage killers of any animal that might be 
unfortunate enough to be in its way, yet like with its more famous 
relative Arctodus (better known as the giant short faced 
bear) first impressions may in this case be deceptive. The post cranial 
skeleton of Agriotherium is that of a large but relatively underpowered 
animal that simply does not seem to have the skeletal framework 
necessary to cope with high stresses, such as those expected to be 
encountered while undergoing extreme physical exertion (i.e. catching 
and subduing struggling prey). The second clue is that Agriotherium has a
 proportionately short snout to that seen in many other bears. The 
advantages of having a short snout are simple, it means that whatever is
 being bitten, is closer to the point of jaw articulation (fulcrum) so 
that greater force can be brought to bear (no pun intended) against it.
These are all features that are common to Arctodus which also has 
isotopic analysis of its bones revealing that it was eating nearly every
 type of animal in its ecosystem, something very unusual for a predator,
 but common for a scavenger. Given the superficial similarity in form 
between Agriotherium and Arctodus, it’s reasonable to speculate that 
Agriotherium may have been a specialised scavenger, a theory that is 
becoming increasingly put forward for Arctodus. Again, the concept is 
very simple, by being bigger than any other predator on the land, 
Agriotherium could in effect bully the smaller predators away from their
 kills. This draws parallels in bear/wolf interaction that is observed 
in the wild even today, where grizzly bears will watch a pack of wolves 
bring down a prey animal, just to charge on in and drive them off after 
they have done all of the work for it. This fits with the surprisingly 
gracile skeleton of a large animal like Agriotherium, since if it was 
letting other predators do the work and the killing for it, why waste 
precious nutrients and calories upon developing and maintaining a 
skeleton stronger than it needed to be?
The idea of Agriotherium being what is termed a ‘hyper-carnivore’ is 
plausible, though it is not certain that Agriotherium only ate meat. 
Like with bears today, Agriotherium may have also supplemented its diet 
with fruits and certain plants, particularly tougher ones that required 
strong jaws. However the scavenger theory does actually fit better with 
Agriotherium in terms of the age of known fossils. Agriotherium first 
appears just after halfway through the Miocene before disappearing at 
the end of the Pliocene. The similar Arctodus however begins to appear 
in the Pliocene before becoming most numerous during the Pleistocene. It
 might be that Agriotherium was one of the first specialised scavenger 
bears but was eventually replaced in the worlds ecosystems by more 
advanced versions that form separate genera, as well as possibly other 
bone crunching animals such as hyena.