What comes to mind when most people think of the Ice Age? It’s often imagined as an endless polar night, a frigid wilderness, and a barren snow-covered landscape stretching to the horizon. In short, it’s not the most hospitable environment for a diverse array of life.

There are contemporary examples of such harsh conditions. We have Arctic deserts, where even lichens and mosses struggle to form a continuous cover. Frozen gravel, boulders, ice as hard as stone, and stone as cold as ice dominate these lands. Even polar bears, the iconic wanderers of the Arctic, are reluctant to venture into these barren territories.

Further south lies the tundra, with its permafrost, vast marshlands, a carpet of mosses, and clusters of shrubs. Here, the soil is thin and infertile, and the landscape is characterized by high humidity. This is the domain of lichens, mosses, low-growing grasses, berries, and dwarf trees. The fauna of the tundra is limited: reindeer, arctic foxes, lemmings, hares, and wolves. There are more birds than mammals, and more fish in the cold, oxygen-rich lakes than there are birds.

This is the state of northern biotopes today, but it wasn’t always so. According to many studies, the modern tundra is a degraded landscape, a remnant of what was once a thriving ecosystem—the mammoth steppe.

Mammoth steppes, or tundra-steppes, emerged during the Pleistocene, in the time of the glacial periods. Their foundation lay in the steppe and forest-steppe landscapes of the Pliocene, and their fauna, known as the mammoth fauna, was a direct descendant of the earlier Hipparion fauna. The mammoth steppes reached their zenith during the last glaciation, 100,000 to 12,000 years ago. This was the most expansive biome on the planet, covering much of the Northern Hemisphere.

The prosperity of the mammoth steppes was due to a fortunate combination of factors:

• Grasses, towering over 2 meters high, covered vast expanses and formed the primary food base. Willows, dwarf birches, and alders grew in the lowlands and along the shores of water bodies, interspersed with patches of larch and spruce woodlands.
• The dry and frosty climate in the periglacial zone turned the mixed vegetation into “standing hay,” easily accessible beneath shallow snow, providing abundant food for numerous herbivores during winter.
• Herds of ungulates trampled the grasses into the ground, helping to form a dense turf that retained nutrients in the soil. They also left behind large quantities of manure, known for being an excellent fertilizer. While permafrost lay just a meter or less beneath the surface, the fertile layer formed with the help of these animals allowed the grasses to recover during the brief and intense northern summers.

In terms of productivity and fauna diversity, the mammoth steppes are often compared to the African savanna. Now, let’s get acquainted with the most prominent and striking inhabitants of this world.

Mammoths are not just a source of valuable fur; they are a whole genus, Mammuthus, within the order Proboscidea, with more than 16 species.

The first mammoths appeared in Africa about 5 million years ago, thriving in the savannas from Cape Town to Ethiopia. Their closest relatives in the genus Elephas, the modern Indian elephant being the sole surviving member, lived alongside them. Both groups migrated into Eurasia via the Middle East, but the ancestors of Indian elephants preferred southern forests, while the mammoths ventured northward. As the Pliocene drew to a close, European rivers were teeming with hippos, and broadleaf forests extended beyond the Arctic Circle.

The oak groves and mixed forests of the temperate zone became home to the southern mammoth (M. meridionalis). Winters grew progressively colder, and the mammoths grew larger. Around 2 million years ago, the southern mammoth was replaced by the steppe mammoth (M. trogontherii), which could grow up to 4.7 meters tall at the shoulder and weigh about 10 tons. This species was likely the first mammoth to have a thick woolly coat, not surprising given its origins in the steppes of southern Siberia. From there, the steppe mammoth spread west to France and east through Alaska to the American prairies. In North America, the steppe mammoth’s descendant, the warm-adapted Columbian mammoth (M. columbi), evolved, while in Siberia, the classic woolly mammoth (M. primigenius) emerged around 800,000 years ago.

Though smaller than its ancestors, standing up to 3.5 meters tall, the woolly mammoth was massive and stocky, with short legs, small ears, and a hump on its back. Males could weigh up to 8 tons, females no more than 4.5 tons, and newborn calves a mere 90 kg.

Now, let’s talk about the fur. The woolly mammoth’s body was covered in a thick fur coat, forming a “skirt” along the sides and belly, similar to that of a yak. Mammoth hair samples appear reddish-brown today, but this is due to the fading of the pigment; in life, the fur was likely dark brown. Genetic analysis shows that occasionally, mammoths had lighter-colored fur.

Another significant attribute of the mammoth was its tusks. They were indeed impressive, reaching a maximum length of 4.2 meters and weighing up to 91 kg. Imagine, when straightened, such a tusk would be taller than the mammoth itself! The tusks were used as weapons in battles between males, for defense against predators, or to dig through snow in search of food. And they needed a lot of food—an adult mammoth consumed at least 180 kg of vegetation daily, mainly sedges and steppe grasses, but also shrubs, flowers, leaves, tree bark, and even moss. Despite their durable enamel plates, the mammoth’s four molars wore down over time, being replaced six times during their lives.

Woolly mammoths went extinct with the end of the Ice Age, around 12,000 years ago, although some island populations survived longer—Wrangel Island’s mammoths, for instance, lived until the second millennium BCE.

The woolly rhinoceros (Coelodonta antiquitatis) is another epic representative of the mammoth fauna that one would love to see in person. Picture this: the tundra, a reddish carpet of dwarf birches, grassy meadows, and a shaggy beast with a long horn on its nose.

The ancestors of the woolly rhinoceros lived 2 million years ago in Asia, north of the Himalayas. From there, they spread across the Tibetan Plateau, China, Mongolia, and reached the shores of Lake Baikal. From this region, the woolly rhinoceros spread across northern Eurasia. In Pleistocene Europe, rhinoceroses were not a rarity—there were already the Merck’s rhinoceros and several species of Elasmotherium. Against this backdrop, the woolly rhinoceros was smaller, less numerous, and more adaptable. Everything changed with the onset of the glaciers 250,000 years ago. The woolly rhinoceros, easily adapting to the cold climate, displaced its competitors and could be found from Britain to Chukotka.

In the tundra-steppe, the woolly rhinoceros was second only to the mammoth in size, growing up to 4.5 meters in length and weighing around 2-3 tons. In its shaggy coat, it must have appeared even larger. A notable feature was the two horns on its snout. The rear horn was small, just 15 cm, but the front one grew to a meter or more. It was flat and broad, used for clearing snow in winter and for combat between males in summer.

Like the mammoth, the woolly rhinoceros fed primarily on grasses, occasionally on conifers and tree leaves. Despite the harsh winters, the snow cover in the tundra-steppe was shallow, 20-25 cm, allowing easy access to vegetation. As the climate warmed, increased humidity led to deeper snow, making movement and access to food difficult. By the end of the Pleistocene, the range of the woolly rhinoceros had shrunk to Transbaikalia and the northern regions of Mongolia and China. It is believed that the woolly rhinoceros went extinct around 14,000 years ago, though some finds suggest they may have survived until 9,000-10,000 years ago.

Elasmotherium, a member of another rhinoceros genus in Eurasia, was less widespread but larger than the woolly rhinoceros. The type species and the longest-surviving one, the Siberian elasmotherium (Elasmotherium sibiricum), is known from finds in the Krasnodar region, along the Volga, near the Southern Urals, and the Irtysh River. Despite its rhinoceros-like appearance, it was not closely related to true rhinoceroses (subfamily Rhinocerotidae). Their evolutionary paths diverged from elasmotheriums (subfamily Elasmotheriinae) as early as the Oligocene, if not earlier. During the last glaciation, Elasmotherium remained the sole species of its once-thriving subfamily. A significant portion of its range was in the steppes of Central Asia, with only a small overlap with the tundra-steppe in the north. Like the woolly rhinoceros, Elasmotherium preferred open spaces and river valleys rich in grasses.

Perhaps its larger size allowed it to compete with the woolly rhinoceros, defending its feeding territories. With a length of 4.5 meters, it stood 2 meters tall at the shoulder and weighed as much as a young mammoth, around 4-5 tons. The horn of Elasmotherium has not been found, and estimates of its length and shape vary greatly. In the Kapova Cave in the Urals, there is a depiction of a humped, massive rhinoceros with a single long horn, interpreted as an image of Elasmotherium. The last Elasmotheriums lived 35,000-38,000 years ago in southern Western Siberia.

The steppe bison is an animal whose prospects for restoration through genetic engineering are among the most realistic, given that it has living descendants. We are, of course, talking about the European bison and the American bison. Both species are descendants of the steppe bison (Bison priscus). In the Pleistocene, herds of these animals roamed the tundra-steppe from Spain to Canada. Large, standing over two meters at the shoulder, with massive foreheads, curved horns, and curly fur, they gathered in enormous herds, followed closely by cave lions, wolves, hyenas, and humans.

Around 120,000 years ago, the European population of steppe bison began interbreeding with other large cattle, the aurochs (Bos primigenius). Despite belonging to different genera, with their evolutionary lines diverging 1 million years ago, this hybridization was successful. Genetic analysis shows that the hybrid bulls were sterile, but the cows produced healthy offspring. Over time, a new species emerged, inheriting the nuclear genome of the steppe bison and the mitochondrial DNA of the aurochs. Today, this species is known as the European bison. In America, the evolution of the steppe bison led to several species, including the giant B. latifrons, with horns spanning up to 2 meters and weighing as much as an SUV. The most successful species, however, was the modern American bison (B. bison), which survives to this day.

The giant deer, a member of the mammoth fauna, was rarely found in the open tundra-steppe. Instead, it preferred the more southern landscapes, with spruce and pine groves, meadows, and thickets of shrubs. The origin of the species is shrouded in mystery, lost in the mountain valleys of Greece, or perhaps the North Caucasus. According to DNA studies, its closest relative is the fallow deer, though the giant deer was significantly larger than the graceful fallow deer.

The giant deer is considered the heaviest of all deer, weighing up to 700 kg, second only to the moose. But its most distinctive feature was not its size (over 2 meters at the shoulder) but its enormous, sprawling antlers, spanning 3.6 meters and weighing 40 kg.

As the Ice Age waned, the giant deer spread across Europe and then expanded its range to Lake Baikal. This expansion was likely influenced by the retreat of the tundra-steppe northward and the spread of forest-steppe areas. Numerous finds have been made in the peat bogs of Ireland, earning the animal its second name—the Irish elk. The extinction of the giant deer is linked to the encroachment of forests, climate change, and human activity. The last of these giants were seen in the northern Black Sea region 7,700 years ago.

No list of the giant herbivores that lived in the tundra-steppe would be complete without this rodent. Unlike the cosmopolitan mammoth, its range was limited to North America, where giant beavers were found in Alaska, Canada, around the Great Lakes, and as far south as Florida.

The giant beaver coexisted with sabertooth cats and short-faced bears, but it was far from easy prey. With a body length of 1.9 meters (and 2.2 meters including the tail), it could weigh over 120 kg. Like modern beavers, it was an excellent swimmer, with webbed feet and a tail elongated like that of a muskrat. The incisors, already large in rodents, grew up to 15 cm long in the giant beaver! Powerful muscles provided a strong bite.

The giant beaver undoubtedly felled trees and fed on their bark, shoots, and aquatic plants. But was it as skilled a builder as modern beavers? Unfortunately, no giant dams have been found, and its brain size suggests that the giant beaver was less intelligent than its modern relatives and not inclined toward complex behavior. Nevertheless, while it may not have built dams, it certainly constructed lodges—remains of which have been found in Ohio’s peat bogs. The last giant beavers disappeared 11,000 years ago, though they may have persisted longer. At the very least, Native American legends have preserved the memory of this fascinating animal.

The dire wolf opens the list of predators in the mammoth prairies of North America. The Canidae family emerged on the plains of America 40 million years ago, and about 6 million years ago, the genus Canis—the “dog” or “wolf” in scientific classification—appeared. At that time, the ancestors of the dire wolf branched off into a separate evolutionary line. Some of them crossed Beringia and spread across the steppes to Africa, eventually evolving into the striped jackal. Those that remained in America faced a different fate. Competing with other canids, they occupied the niche of hunters of large prey. Thus arose the largest canids (excluding certain dog breeds)—the dire wolves (Aenocyon dirus).

On average, male dire wolves grew up to 1.7 meters in length and weighed around 68 kg. Only the largest specimens of regular wolves from Siberia and Canada can boast similar sizes. Larger skeletons have been found in the La Brea tar pits, and biomechanical analysis suggests that the maximum weight of a dire wolf could reach 110 kg, double that of a typical wolf.

The northern boundary of the dire wolf’s range was in the periglacial zone, slightly south of the current US-Canada border. Here, bison, horses, camels, and pronghorns provided the main prey for the dire wolf. They weren’t above scavenging, as evidenced by the characteristic wear on their teeth. Like gray wolves, dire wolves were social animals. At the head of the pack was an alpha pair, male and female, with the rest of the pack consisting of their offspring of various ages. Coordinated actions allowed them to kill animals weighing 300-600 kg, making the dire wolf a serious competitor to Smilodon and the American lion. The dire wolf went extinct 13,000 years ago due to climate change that reduced their food supply and the spread of the more adaptable gray wolf.

The cave hyena was a large subspecies of the spotted hyena, inhabiting northern Eurasia from 500,000 to 11,000 years ago. It was distinguished by its larger size—the cave hyena stood up to a meter tall at the shoulder and could weigh over 100 kg, almost twice the average weight of the spotted hyena. The cave hyena’s range covered a vast territory, from the Iberian Peninsula to the Far East. These predators preferred steppe landscapes with natural shelters—groups of rocks, grottos, and caves.

Like modern hyenas, the cave hyenas preferred hunting over scavenging. The bones of animals found in their dens indicate that hoofed animals were their primary source of food. Horses made up the bulk of their diet, followed by bison and woolly rhinoceroses. Less frequently, they hunted deer, saiga, wolves, cave lions, and bears. Relations with humans were complex. Neanderthals and cave hyenas alternately occupied the same caves, suggesting some level of competition. Cave paintings in France depict hyenas, and hyena bones show signs of human processing. However, human skeletons gnawed by hyenas are more commonly found.

The Homotherium was a sabertooth cat that roamed the northern reaches of the tundra-steppe, from Europe to Alaska. In America, south of Canada, its remains are rarely found, with the southernmost being in Venezuela. It appears that in these territories, it was outcompeted by the larger Smilodon, but in the periglacial zone, it was the only sabertooth cat.

Homotherium could weigh up to 190 kg and stand 1.1 meters tall at the shoulder, comparable to African lions. It had somewhat hyena-like body proportions, with long front legs and shorter back legs. Its tail was short, like a lynx’s or bear’s. It was likely a good runner, as evidenced by its non-retractable claws, relatively thin limbs, a nasal opening similar to that of a cheetah, and a complex and large visual cortex. Modern data suggests that Homotherium lived in prides, which facilitated hunting and the protection of offspring.

Like other sabertooth cats, Homotherium specialized in large and very large prey. Remarkable finds have been made in the Friesenhahn Cave in Texas, where more than 30 skeletons of Homotherium of various ages were discovered, along with the remains of 400 young mammoths, whose carcasses had been dragged into the den and consumed there.

The cave lion was another “African” species widely spread in the tundra-steppes of the Northern Hemisphere. For a long time, it was considered a subspecies of the African lion, but genetic studies have disproven this. The first lions appeared 1.7-1.2 million years ago in Tanzania. In Europe, their descendant, the Mosbach lion (Panthera leo fossilis), appeared about 700,000 years ago and spread eastward to southern Western Siberia. The advance of the glaciers led to the emergence of the cold-adapted cave lion (Panthera leo spelae), which crossed Beringia into America, where it evolved into the American lion (Panthera leo atrox).

The cave lion was larger than modern African lions, though it did not reach the size of the Mosbach or American lions. With a body length (excluding the tail) of over 2.5 meters and a weight of up to 330 kg, the cave lion was one of the key predators in the tundra-steppes. A significant portion of its prey consisted of reindeer and cave bears, followed by horses and musk oxen. Occasionally, remains of young woolly rhinoceroses and mammoths killed by cave lions have been found. This hunting specialization likely reduced competition with other predators but may have played a role in the cave lion’s extinction.

The cave lion is a frequent character in artistic depictions of the Stone Age. Primitive hunters left us impressive images of these animals. Interestingly, these images reveal that the cave lion lacked a mane. In addition to bones and drawings, four lion cubs have been found in the permafrost of Yakutia.

The cave bear, along with the mammoth and woolly rhinoceros, can be considered an iconic figure of the Ice Age. Over 1 million years ago, it shared a common ancestor with the brown bear, but their evolutionary paths diverged. The cave bear was characterized by a significant increase in size and a predominance of plant-based food in its diet. It could weigh twice as much as its brown bear relative, and standing on its hind legs, it reached a height of three meters. Comparable sizes are achieved today by the brown bears of Kodiak Island, easily surpassing 500 kg.

The wear on its teeth and the loss of premolars indicate a greater reliance on plant food than the brown bear. Succulent grasses and roots likely made up a significant part of its diet. However, the cave bear should not be considered a gentle giant. It could certainly take down a deer or young bison if given the chance, though it also often fell prey to predators like the cave lion. It seems that cave lions would search out bear dens in the dark of caves and attack them during hibernation.

Interestingly, the cave bear was the most genuine “spelunker” among the “cave” animals of the Pleistocene. Its claw marks and skeletons have been found deep within the earth, in underground grottoes that even humans find challenging to access. It was in these caves that the bear made its dens, ultimately making the species vulnerable to humans. In the Stone Age, humans hunted cave bears and worshipped them. Neanderthals kept bear skulls as trophies. Cave bear depictions are known from cave paintings, and it’s possible that the bear totemic cults known among the taiga peoples have their origins in the Paleolithic era.

Arctodus, or the giant short-faced bear (Arctodus simus), concludes our list as the largest predator among the mammoth fauna. As a member of the subfamily Tremarctinae, it is only distantly related to other bears. Its ancestors lived in North America as far back as the Miocene. When the Isthmus of Panama formed 2.5 million years ago, some of them migrated to conquer South America, resulting in the spectacled bear (Tremarctos ornatus), the only bear species on the continent and the most herbivorous member of the bear family. Those that remained in the north turned into true giants 800,000 years ago.

The shoulder height of this monster ranged from 1.5 to 2 meters, taller than a woolly rhinoceros. Claw marks in Missouri’s Riverbluff Cave, found at a height of 4.6 meters, suggest that it could stand over 3.7 meters tall on its hind legs. The bear’s weight exceeded 950 kg, and in northern populations, there may have been giants weighing up to 1,400 kg, making Arctodus one of the largest terrestrial carnivorous mammals in history.

Just how dangerous was this beast? Current understanding suggests that it moved, like all bears, at a slow, deliberate pace but with enormous endurance. When attacking, Arctodus could reach speeds of up to 40 km/h, quickly overtaking and knocking down its prey. This allowed it to kill bison, mastodons, and young mammoths. However, current thinking suggests that Arctodus was not a superpredator like the polar bear. Its feeding behavior was flexible, similar to that of the brown bear. Northern populations were largely carnivorous, while bears living further south consumed a significant amount of plant material.

The inhabitants of the mammoth steppes with whom we’ve become acquainted are part of the Pleistocene megafauna. They are striking, unusual, and capture people’s attention. But we mustn’t forget that alongside mammoths and Arctodus lived “smaller” species, most of which still exist today. Reindeer, musk oxen, saiga antelope, yaks, Bactrian camels, and horses are still with us, though they inhabit different landscapes. Predators such as wolves, arctic foxes, red foxes, lynxes, stoats, martens, and wolverines also remain. Lemmings, hares, numerous birds, fish, and insects still share the planet with us. The plants that provided food for the animals of the tundra-steppe have also survived.

This means we have a chance to restore this unique landscape. In fact, a similar experiment is already underway in Yakutia. The Pleistocene Park, founded by ecologist Sergey Zimov in 1997, now houses Yakutian horses, American bison, European bison, musk oxen, red deer, moose, and reindeer. And, incredibly, the ancient ecosystem is beginning to recover, forming patches of a new tundra-steppe.

At the same time, research on the DNA of extinct members of the northern megafauna is ongoing. Ambitious projects by Japanese and Korean scientists to clone the mammoth are just the tip of the iceberg. Perhaps the cave bear, wild horse, or steppe bison will prove to be more suitable candidates for the first resurrected animal. Over time, Pleistocene Park might transform into a fully-fledged mammoth steppe, where herds of shaggy giants roam once again.