Rabbits in the Ice Ages

The rabbit (Oryctolagus cuniculus) is one of the more familiar mammal species to most people in the developed world, not least because it has been partially domesticated. Moreover, wild and feral rabbits are found across wide parts of the world, living on every continent except Antarctica. They are adaptable animals, able to survive in a wide range of different habitats, from hot semidesert to forests and bleak moorland. So much so, in fact, that, in most parts of the world, they are a pest.

The rabbit is widespread because we humans have spread it, following the initial domestication event in France, no later than 800 AD. This was originally for meat and fur, with pet breeds appearing only from the late 18th century. Rabbits are now found, for example, on Middleton Island, a chilly speck of land 130 km (80 miles) off the south coast of Alaska, and on the Kerguelen archipelago in the Indian Ocean, which is about as remote a place as it's possible to get.

How the Lemming Got its Coat

The Ice Ages were, without doubt, the most dramatic natural climatic changes on Earth in the last few million years. The last one was particularly severe, with vast ice sheets covering much of northern Europe and northern North America. This, naturally enough, forced many species of animals in the Northern Hemisphere to move south. Even those well-suited to the cold, such as reindeer, musk oxen, and woolly mammoths, would have had to avoid the barren ice sheets, even if they were happy in the broad tundra belt to the south.

In Europe, in particular, there is only so far south you can go before hitting the coastline. This meant that many animals were forced into small areas, some of which may still have been marginal habitat for them, to avoid extinction. These areas are called "refugia", and their small size and isolation were a driver for evolutionary change. Sometimes populations were split apart for so long that they became separate species and, for example, we can date many species of northern birds to this time, even though, for them, the Mediterranean would not have been an issue. 

When White-footed Mice Invaded Michigan

When I'm describing a particular species of mammal on this blog, one of the first things I usually mention is where in the world it lives wild. Clearly, there are differences in the fauna of different continents or specific islands. Cougars, black bears, pronghorn antelope, and coyotes are all common enough in the US, but don't live in Europe. We have Eurasian and Iberian lynx, but not bobcats and Canada lynx. There are bison in Europe, but they're a different species than the American sort, and we only have wild raccoons because some were deliberately released in Germany in 1934. 

Clearly, the existence of the Atlantic Ocean is not to be sniffed at. However, this applies on a smaller scale, too, where some physical barrier that's some way short of an ocean, but is still significant, prevents an animal from advancing further across its home continent than it might like. 

And then there's the fact that animals have particular requirements as to the climate and vegetation of their native areas - even if the vegetation is only affecting the prey animals that they themselves need to feed on. The difference between this and the physical geography of rivers, mountains, and so on, is that it's changing over a much faster timescale, especially in recent decades. Animals may be forced to move to new areas, which can be a problem if the physical geography prevents them from doing so and can be a problem for other reasons even if it doesn't.

Pleistocene (Pt 15): Ice Age Down Under

Procoptodon

The island continent of Australia has today what is probably the strangest mammalian fauna of any continent. Yet it is also the continent that has, perhaps, suffered the greatest number of mammalian extinctions over the last 50,000 years or so. Many of those are relatively recent, or at least after the end of the last Ice Age around 10,000 BC. But even compared with the Australia of, say, the 18th century, the wildlife of Ice Age Australia looked pretty odd.

In climactic terms, Australia didn't suffer too badly from the Ice Ages. It's too close to the equator to have had ice sheets get anywhere near it, although doubtless there was rather more snow on the mountains. (Although perhaps not too much - even today, Australia is the only continent to lack glaciers). Then, as today, much of the continent consisted of desert, and the bits that weren't were mostly arid grassland, albeit with denser woodlands around the eastern and northern coasts.

Pleistocene (Pt 14): Ice Age Africa

Giant warthog

In the earlier parts of this series, when I described the animals of Ice Age Europe, I said that the continent, at that time, had a feel reminiscent of present-day Africa. But, if that's so, what was Africa like during the Ice Ages?

The answer is, perhaps disappointingly, "not that different to how it is now". Indeed, of all the settled continents, it's probably Africa that has changed the least since the Ice Ages. You might think that this has something to do with Africa being close to the equator; in particular, that it's too close for whopping great sheets of ice to have rolled across the countryside.

Which they didn't, so that much is true. Indeed, the southern hemisphere in general had far less ice cover than Europe, Asia, and North America. That's due mainly to the way that the continents happen to be arranged, with glacial ice sheets only being able to get as far as southern South America (there are fjords in places like Tierra del Fuego). Presumably, sea ice extended much further across the Southern Ocean than it does now, but that would have little effect on land-based animals.

But that's not to say that Africa, or Australia, were unaffected by the Ice Ages. Africa was colder than it is today, and, to begin with at least, rather drier, too. The Sahara and Kalahari deserts were more extensive than they are today, with the semi-desert belt of the Sahel being quite a way south of its present position, running through what are now fairly lush countries such as Guinea, Nigeria, and northern Kenya.

Pleistocene (Pt 10): Of Armadillos and Bison

Glyptotherium

When the continents of North and South America collided, shortly before the dawn of the Pleistocene, a range of creatures wandered across the newly formed isthmus of Panama. Most were heading south, giving us such "normal"-looking South American animals as jaguars, ocelots, and grey foxes. The ones heading north were a lot stranger, and many of them didn't do so well in the long term. These include such wonderfully weird creatures as the giant ground sloths, which managed to get as far north as modern Canada before dying out.

But they weren't alone, and not necessarily any stranger than some of their fellow immigrants. The same event that saw the arrival of the ground sloths also saw the coming of the armadillos. Today, there is only one species of armadillo in the United States, and only two outside of South America (the other gets no further north than southern Mexico). But, back during the Pleistocene, there were others, and some of them were not quite what we'd expect today.

Pleistocene (Pt 8): Mammoths v. Mastodons

American mastodons

The arrival of the first mammoths in North America was a significant turning point in the development of the local wildlife. It's so important that this date, 1.9 million years ago, marks the beginning of the first of just two 'land mammal stages' that define North American wildlife during the Ice Ages. It used to also mark the beginning of the Pleistocene itself, but for various reasons, that's now been shifted a little further back.

The mammoths in question arrived from Asia, crossing over the Bering land bridge, the recurring appearance and disappearance of which greatly influenced North American wildlife during this time. They were southern mammoths (Mammuthus meridionalis), the dominant species of mammoth in Asia at the time, but they quickly evolved into a home-grown American animal: the Columbian mammoth (Mammuthus columbi).

It used to be thought that, even ignoring any late-surviving southern mammoths, there were at least two different species of mammoth living in North America in the early to mid Pleistocene. We're now pretty confident that they're all just examples of Columbian mammoth. Nonetheless, you will often see references to the "Imperial mammoth" (Mammuthus imperator). Perhaps the biggest elephant that has ever lived - they were about thirteen feet tall at the shoulder - these were probably just really big Columbian mammoths. Not that that's anything to sneeze at, mind you.

Pleistocene (Pt 7): Meanwhile, Across the Atlantic...

Columbian mammoth
(It's likely that the real animal was hairier than the one in
this reconstruction, but it shows the tusks effectively)

Over the last five parts of this series I have described the history of Pleistocene Europe, describing some of the ways that the animal life of the continent changed over those thousands of millennia, and looking at a few particular animals in more detail. But one doesn't need a degree in zoology to notice that today, the wildlife of North America, for example, is different to that of Europe. North America has coyotes, raccoons, cougars, armadillos, and pronghorn antelope, to name just a few animals that are simply absent in Europe. (Or, in the case of raccoons, were absent until somebody made the mistake of releasing some of the furry nuisances in 1930s Germany).

It's hardly surprising that that continent also had different wildlife during the Pleistocene. One notable difference, for instance, is that we humans weren't there. Obviously, neither Columbus nor Leif Ericsson were genuinely the first person to discover America. But even whichever long-lost group of Native Americans was actually the first to discover the great western continent, they did so long, long, after the first Europeans discovered Europe. While Europe had at least some species of human inhabiting it for about two-thirds of the Pleistocene, nobody reached America until the epoch was all but over.

Pleistocene (Pt 6): Lions, Hyenas, and Bears, oh my

Cave lions
(original artist unknown, c. 33,000 BC)

The iconic Ice Age carnivore is, of course, the sabretooth cat. However, as I mentioned when discussing the Mid Pleistocene, sabretooth cats when extinct in Europe well before the Last Ice Age. (Well, maybe not... but at any rate, they seem to have been very rare). So what were the dominant predators in Europe during that final, and most extreme, period of intense cold?

It probably depends on how small a predator you're willing to count. Animals like weasels, stoats, and shrews were presumably very numerous, even if their skeletons are often too delicate to preserve well. Moving up the scale, the most numerous carnivoran fossils from the time belong to foxes. Both the modern red fox and the Arctic fox were widespread in central Europe at the time, although the latter was presumably more comfortable.

Another modern animal that we wouldn't be surprised to find among the snowy forests and open tundra was the grey wolf. Indeed, grey wolves may well have originated in Europe from an American ancestor, before heading back over the Bering land bridge. If so, they were probably quite quick about it, since they are found across the Arctic on both sides of the Pacific from an early date.

All of these are creatures that are still found in Europe today, although not necessarily in large numbers - wolves are more common in the wilds of Asia and North America than they are in densely settled Europe. But I've been harping on through this series about how "African" European wildlife looked at the time, and that's as true of the big carnivores as it is of herbivores such as elephants/mammoths and rhinos. One of the clearest instances of this is the presence in Pleistocene Europe of lions.

Pleistocene (Pt 3): Ice Ages and Interglacials

Life-size reconstruction of a steppe mammoth
(compared with a 3-year old human)

When the Pleistocene began, Europe's climate was much the same as it is today, and the general shape of the continent would also have been instantly recognisable from space. The animals however, were different, many of them being ones we would now associate with Africa - rhinos, elephants, hyenas, and cheetahs, among others. In part 2, I described how that began to change 1.8 million years ago (which, incidentally, was once defined as the beginning of the Pleistocene - see part 1 for why that changed).

This was a time of cooler weather, as the Ice Ages began to dawn. Forests retreated in the face of advancing tundra, and musk oxen, bison, and (strangely) European hippos began to make their appearance. The cold snap was prolonged, and, so far as we can tell, the fauna of Europe remained relatively stable for the next 600,000 years. That's still a very long time - if we go back to my analogy where we get just one minute to watch the events of a decade, with the whole of written history thereby spread out into a nine hour spectacular, this phase of European history would last a full six weeks.

1.2 million years ago, half way through the Pleistocene, the climate changed again, and mammals (and other animals) were forced to adapt. However, the change wasn't towards yet colder weather, but back towards a warmer, more pleasant climate. The forests grew back, with all their dense undergrowth in attendance, and the harsh steppe-lands retreated into the north. As had been the case at the dawn of the Pleistocene, European weather would have been much as it is now.

Pleistocene (Pt 2): Europe at the Dawn of the Ice Ages

Pachycrocuta brevirostris, a European hyena

The Pleistocene is the time of the Ice Ages, when great ice sheets rolled across much of the northern hemisphere. Nothing much lived on the ice sheets themselves, just as there is very little today in the heart of Greenland. But, as we've seen, not only were their wide bands of tundra and pine forest reaching across much of today's 'western' world (and, of course, a fair chunk of the Orient), but the ice ages weren't continuous; there were many warm gaps between them.

The mammals of the Pleistocene include what are surely the most familiar fossil mammals to most people, the ones we generally think of when we think of 'after the dinosaurs'. For this was the time of the mammoths and sabretooths. They're familiar to us because, aside from the tiny sliver of warm weather we currently live in, the Pleistocene is the most recent, and therefore the best preserved and the most easily analysed, of all the epochs of the Age of Mammals. It's been the setting for a number of films, books, and TV series - to name just two, the Ice Age cartoons, and the Earth's Children series.

Yet, when we think seriously about Pleistocene animals, there are a couple of important points to bear in mind.

Age of Mammals: the Pleistocene (Pt 1)

A scene from northern Spain

The "Age of Mammals" is the informal name for the Cenozoic era, the 65 million year slice of Earth's history from the extinction of the dinosaurs to the present day. It is so named because mammals have been the dominant large, land-dwelling animals throughout the era.

Of course, "large, land-dwelling" is something of an arbitrary qualification, and one more rooted in the natural prejudices of our own species than in an actual reflection of Earth's biodiversity. The most numerous animals throughout the era, and, for that matter, through the Age of Reptiles that preceded it, would have been insects. But, unless you're standing in the middle of a swarm of midges, most people don't notice insects in the same way they would notice, say, a herd of antelope, or a prowling tiger. Mammals aren't even the most numerous vertebrates today, and, by sheer species count, it's the fish that are dominant, and some of those are pretty big.

Even on land, in terms of number of species, mammals are the least numerous of the four vertebrate classes - birds come in first, and reptiles still hold on to second place, followed by amphibians. Of course, most of those reptiles are small lizards, and birds are also generally quite small. Even so, there are ostriches, crocodiles, and anacondas, among others, and, in fairness, most mammal species are mouse-sized. So there's a reasonable case that what this should really be is the Age of Birds. But I, for one, am going to stick with the standard term.