Indricotheriinae is a subfamily of Hyracodontidae, a group of long-limbed, hornless rhinoceroses that evolved in the Eocene epoch and continued through to the early Miocene. The earlier hyracodontid species, such as Hyracodon were modest-sized, fast-running, lightly built animals with little similarity to modern rhinos. However, during the late Eocene and early Oligocene, the indricotherines evolved, and quickly grew to huge sizes. They flourished in the rainforests of a coastal region that became Kazakhstan, Pakistan, and southwest China, and lived further inland throughout central Asia as well.

The indricotherines reached the peak of their evolution from the middle Oligocene through to the early Miocene, where they had become truly gigantic animals, represented by the specialized genera Indricotherium and Paraceratherium (these genera may be synonymous, but several distinct species probably remain valid). These were the largest land mammals that ever lived, equaling the medium-sized sauropod dinosaurs in size. However, they remained confined to central Asia, which at the time was part of a large, lush lowland region. The collision with the Indian subcontinent and the Himalayan uplift led to global cooling, desertification, and the disappearance of forest habitats, which resulted in the extinction of these giant ungulates. Certain unidentified species may have lived in Mongolia, and many other places in Russia.


No evidence has turned up that the giant short-faced bear, the sabertooth cat,or any of the other extinct big beasts shown in this painting
(save for the mastodon) were ever hunted by early Americans

One of archaeology's enduring mysteries is why animals like giant sloths, mammoths, and giant beavers died out suddenly at the end of the last Ice Age. Current thinking regarding these Pleistocene and Holocene extinctions promotes climate change, human hunting, and trans-species disease as the main culprits. These competing theories can be thought of as "chill-kill-and-ill."

Ecologist and "kill theorist" Paul S. Martin argues that humans possessed both motivation and opportunity to hunt the mega beasts into extinction. Walking along the railroad tracks at Dent, Colorado, archaeologist Gary Haynes agrees, citing as evidence 15 slaughtered mammoths discovered there 60 years ago. Period footage of the original Dent site excavation is augmented by a computer-generated attack of three dozen Paleoindian hunters upon a bewildered herd of mammoths.

Paleontologist Ross MacPhee's promotes the "ill theory," which suggests the mega beasts were taken out by disease--plagues of rabies, bubonic plague or perhaps anthrax--introduced by migrating human newcomers and their "cloud of microbes." On the theory that "where animals get sociable, they get sick," MacPhee examines a "mega beast freeway" at Monster Beach, Bahia Blanca, south of Buenos Aires, where the fossilized footprints of giant ground sloths and other Pleistocene animals are preserved.

Veterinarian Sarah Cleveland suggests that the devastating spread of canine distemper in modern-day East Africa might provide a model of how such catastrophic diseases operated in the past. Showing killer diseases on television is grisly business, and the viewer is subjected to seemingly endless, if dramatic, footage of wild dogs and a lion writhing in their death throes.

Paleontologist Russell Graham points to climate as the main suspect, making the case that a sudden cold snap at the end of the Ice Age triggered the massive die-off. The program concludes, perhaps predictably, with a compromise, suggesting that a nuanced interplay of kill-chill-and-ill might best explain the mega animal extinctions: "In short, what killed the mega beasts was progress."

The Perils of Being Huge: Why Large Creatures Go Extinct
By Bjorn Carey
LiveScience Staff Writer
July 18, 2006

Once upon a time, a 2-ton wombat lumbered across the Australian Outback. Around the same time, mammoths and saber-toothed tigers had the California coastline all to themselves.

Millions of years before any of these animals existed, Tyrannosaurus rex and other colossal dinosaurs ruled the world.

These and some of the other largest and most fantastic creatures ever to walk the planet are long gone, victims of mass extinctions of large beasts. And for reasons poorly understood, often the animals to fill the voids were tiny by comparison.

Predisposed to extinction

Scientists generally accept that a giant asteroid slammed into the Gulf of Mexico some 65 million years ago, setting off a chain of catastrophic events that ultimately led to the extinction of dinosaurs. Whether or not an asteroid is to blame, the so-called KT boundary in the in fossil record displays a mass extinction of dinosaurs and other large animals around the world.

Small scavenging mammals and birds survived the event, and scientists can't say for sure why dinosaurs did not.

Since bigger beasts couldn't take shelter in small protected burrows, perhaps they were done in by fierce environmental conditions. Or maybe with so many plants dying off, big herbivores simply had nothing to eat, and as they died out, so did the big carnivores.

Or perhaps with all the stress, dinosaurs simply couldn't reproduce quickly enough to keep up with sexually nimble mammals and were soon outnumbered.

"If a disease or climate impact is severe enough to kill off most of the young in a generation, it will take a very long time to replace them," zoologist Alex Greenwood of Old Dominion University told LiveScience. "Smaller mammals like rodents, for example, would not be as severely affected since they have multiple young and a very short birth cycle."

Along with mammals, turtles and crocodiles, which can lay hundreds of eggs at a time, managed to survive the mass extinction. Also, because they could take shelter in water, these reptiles probably weren't competing with land mammals for resources.

A mammoth lesson

Then mammals got bigger. And eventually they paid the price.

Several mammoths and other big mammals died off during the Pleistocene/Holocene extinction event, which started around 50,000 years ago and continued through the end of the last major ice age about 10,000 years ago.

Today's large mammals— often with small populations, long gestation periods and late weaning ages—are similarly predisposed to sudden mass extinction, scientists say. For big beasts, taking care of offspring is typically a time sink and an energy drain, and the whole setup makes the young highly susceptible to predation.

Large mammals are also slow to reach sexual maturity, and mortality rates before that age are generally high. The slow pace at which new individuals are introduced to the population presents an obvious challenge to enduring tough times when population numbers drop precariously low.

Super predators

Mass extinctions occur with surprisingly regularity over the long haul. During the last 250 million years, there's been a big die-off roughly every 26 million years.

Adam Lipowski, a researcher at Adam Mickiewicz University in Poland, suggests the extinctions might sometimes be driven not by climate change or impacts from space, but by the emergence of super predators.

In 2005, Lipowski developed a computer simulation representing a population of many species competing for food and living space. Much of the time, "medium efficiency" predators prowled the virtual world and their numbers fluctuated only slightly in response to changes in prey population size.

But every so often, mutations would lead to the evolution of a super predator that quickly devours an entire prey population, which in turn leads to its own extinction.

The critters that survived the "predatory apocalypse" gradually mutated to fill new ecological niches, and the cycle began anew.

Look in the mirror

Humans could be considered today's super predators.

We've been implicated in the extinctions of mammoth, various saber tooth cats, and giant apes, as well as more recent extinctions of dodos, bonobos, and wild horses. Whether human hunting pressure, climate change or other factors were responsible remains controversial, however.

Meanwhile some whale species, among the biggest mammals that ever lived, have seen their populations drop to mere thousands in response to over-hunting and environmental pressures. Scientists estimate only about 300 North Atlantic right whales exist, with much of the attrition attributed to collisions with large shipping vessels and noisy ports causing navigational confusion.

Illegal poaching and habitat reduction have cut African elephant numbers in half, to 600,000, in the last 30 years. Scientists have determined that African lions'historic distribution has shrunk 82 percent in the last few decades and the wild population is estimated in only the low tens of thousands. And although Florida manatee numbers have nearly tripled in the last 15 years, the population hovers around just 3,000.

Humans can also initiate extinctions inadvertently.

Between 55,000 and 45,000 years ago, humans first set foot on Australia. At the time, large emu-like terrestrial birds and oversized wombats roamed the continent. But when humans started setting fires to clear land or flush prey from bushes, they also stripped the land of many of the plants the large animals favored.

Many of the smaller animals that adapted to eat the remaining plants survive today, while the two-ton wombat is no more.

Carnivores Can't Get Much Bigger
By Robin Lloyd
Senior Editor
January 16, 2007

Lions and tigers and bears are about as terrifying, size-wise, as it’s going to get in the wild world of mammals, new research shows.

Ecologists at the Zoological Society of London modeled the energy budgets of land-dwelling carnivores and arrived at a 1-ton limit as the maximum sustainable mass for these meat-eating mammals.

After that, there is a huge pay-off for each kill made by a large predator, but it takes too much lumbering energy for the behemoths to hunt the vast quantities of meat required to carry on across evolutionary time, said Chris Carbone, lead author of the new research published in the journal PLoS Biology.

At least four terrestrial, carnivorous mammals that Carbone puts near the 1-ton mark have gone extinct: the short-faced bear; North American lion; South American sabercat; and Megistotherium osteothlastes (a wolf-like carnivore that lived 25 million years ago in Africa).

Today’s large carnivorous land mammals, including the record-setting polar bears—which usually weigh around half a ton but can grow to weigh nearly a ton, are threatened in part because of the energy intake-and-expenditure equation, Carbone told LiveScience.

“By being a carnivore that specializes on hunting big prey, it potentially can make you vulnerable to the process of extinction,” he said. “It’s more costly to feed on these larger prey, so it’s more energy demanding. And you’re more reliant on prey. If you can’t meet those energy costs, you’re no longer viable over time.”

Hunting strategies

Carbone’s model also explains some of the hunting strategies used by carnivorous mammals and how they vary depending upon the size of the carnivore and whether they tend to prey on invertebrates (such as insects) and smaller animals or larger vertebrates, such as deer, antelopes and their relatives.

Carnivores travel over some of the widest distances of all land-based mammals for their size. The long travels, plus chasing, pulling down and pulling apart of prey costs a lot of energy.

For carnivores that feed on rodents, small reptiles or insects, most of the energy cost comes simply from getting to and from prey, Carbone said. “They might chase and eat it, but it’s dispatched easily.”

Lolling about and naps are ways big carnivores make up for the energy costs, Carbone and his co-authors surmise in the research.

Mid-size carnivores around the size of a lynx, jackal or termite-eating aardwolf, a hyena-like carnivore that lives in Africa, have a cross-over advantage energetically, Carbone said. They have low metabolic rates that allow them to conserve energy and can hunt both small and large prey.

For instance, Eurasian lynxes can feed happily on rabbits and smaller prey, but due to their size, they can also take down medium-sized antelopes. In the absence of large prey, such animals can always resort to rodent-killing, Carbone said.

The difficulty of catching and killing larger animals costs twice as much energy (even though the payoff is greater too). But massive, large-prey predators have no small-prey options—those meals are too small to be worthwhile. Massive mammals must hunt the big prey that require huge amounts of energy to locate, catch and take down. “There is no where else to go, and then you’re kind of stuck,” Carbone said.

Animal 'ground rules'

Paleobiologists will find Carbone's model useful, said John A. Finarelli, a University of Chicago paleontologist who studies all dog-like carnivores. In the past, some scientists have looked at the minimum mass limits for warm-blooded animals, and even stated that North American mammals never approached maximum mass limit.

"However, in light of (Carbone's) findings, it could very well be the case that carnivorous mammals do not play by the same set of rules that other mammals do," Finarelli said.

The largest known land-dwelling carnivores of all time were lower-metabolism dinosaurs such as Spinosaurus (about 8 tons), Giganotosaurus (about 8 tons) and Tyrannosaurus (about 6 tons).

And then there was the 15-ton Indricothere—but that extinct mammal was an herbivore.

“Carnivores, because of the cost of hunting, can never achieve the sizes and intake rate of prey of the largest herbivores,” Carbone said. “It’s costly to be a big carnivore. The weaponry required to take down big prey adds to cost in terms of movement and maintenance. Ultimately that limits your ability to hunt, which again limits your size.”