A very brief introduction to mammoths

I was in Los Angeles when I first saw him. I stood in his shadow as he towered above me. I didn’t feel fear. Or panic. I felt awe despite being dwarfed. Light bounces off each bone, highlighting the curves, the detail, the sheer size. A gigantic skeleton; the internal frame holding up a body of immense proportions. I recognised each bone as I would old friends: a femur at the top of a back leg, vertebrae running along the back of the animal, ribs protecting what would have been the delicate organs beneath. Yet despite him and I being practically the same bone for bone, his were enormous, thicker, stockier, stronger. What’s more, he had two enormous tusks protruding from his face, which curve dramatically outwards and in on themselves. This magnificent beast was the Columbian Mammoth. A goliath from the past.

How did this giant find itself in Los Angeles when elephants alive today are found in Africa and Asia? Was it related to the Woolly Mammoth? Was it hairy? Did humans see these animals? What is a mammoth?

One enormous skeleton. A mountain of questions.

To find the answers we will have to travel back in time and space. Back to Africa. Back to the very beginning of this iconic group of animals.

Yours truly standing under a composite skeleton of a magnificent Columbian Mammoth. The specimen is on display at the Page Museum, La Brea. This incredible museum and active excavation site is sat right in the middle of busy LA. (Photo by random museum visitor)

Mammoths belong to the Order Proboscidea, which has its roots back in the African Paleocene around 60 million years ago. (We, for example, are classified in the Order of Primates. We share this Order with the other primates: gorillas, chimpanzees, monkeys, gibbons, lemurs and lorises) The Proboscidea was pretty spectacular and included many weird species, such as the odd pig-hippo creature, Moeritherium and one of the largest mammals to have walked the Earth, Zygolophodon.

This Order was once astonishingly diverse with 13 different Families holding dozens of different species that migrated into Europe, Asia and the Americas. Just three species of this once successful group survive today; the African Savannah Elephant (Loxodonta africana), the African Forest Elephant (Loxodonta cyclotis) and the Asian Elephant (Elephas maximus), all belonging to the last surviving Family of the glorious Proboscidea; the Elephantidae.

The variety of proboscideans (Artworks by Vladimir Nikolov.
Editing and digital work by Docho Dochev.) (Image from here)

Originating in Africa just around 5 million years ago the Elephantidae has included a huge number of species, from the ‘first elephant’, Primelephas, to the giant extinct straight tusked elephant, Palaeoloxodon. It is special because it is the last surviving Family of a once great and diverse group. It is also special because it includes the mammoths!

The word ‘mammoth’ has slipped into our everyday language as an adjective meaning enormous, massive or huge. (I do quite like it when I hear a friend or colleague describing something as mammoth without realising it.) It has also become synonymous to describe any large hairy ‘elephant’. Forgive me for a moment while I get a little bit passionate about what is and what is not a mammoth.

Mammoths are any species belonging to Mammuthus. Not other large Probosidea or other elephant relatives. Mastodons were not mammoths: they lived on a separate branch to mammoths for around 28 million years. (Mastodons belong to the genus Mammut, confusing, but still not a mammoth.) The freaky Gomphotheres were not mammoths. Nor were the huge Straight Tusked Elephants. Only mammoths were mammoths. These other large beasts may superficially look like mammoths, but they are not. They are, however, wonderful examples of convergent evolution, where similar features evolve in different animals what live similar lifestyles: a classic example is the beautifully streamlined bodies of Ichthyosaurs (marine reptiles) and dolphins (mammals).

The First Mammoth
Now you know what a mammoth is not. What then makes a mammoth: how do we know what species belong to Mammuthus? One diagnostic features of a mammoth is the inward curving tusks, which you saw in the magnificent Columbian Mammoth we met earlier. The skulls of mammoths were flatter and higher than their relatives, which went arm in arm with more highly crowned teeth for a change in diet. We see the beginning of these features back in the Pliocene around 5 million years ago. Indian elephants and mammoths shared a common ancestor and lolloped their separate ways sometime about 5 million years ago. One of these populations evolved into first true mammoth (Mammuthus subplanifrons). Fossils have been recovered in Eastern and Southern Africa, with associated fossils suggesting a warm, tropical environment. From this warm loving (hairless) giant stomped a whole parade of different species!

The Frist Mammoth (Mammuthus subplanifrons), evolved in Africa around 5 million years ago. (Image by Twilight Beasts)

The African Mammoth
M. subplanifrons, the First Mammoth, disappears around 3 million years ago, around the same time the African Mammoth (M. africanavus) appears. One small population of the First Mammoth evolved into the African Mammoth, while shortly after the rest of the first founding species vanished. Relatively smaller than more recent mammoths, the African Mammoth was distributed across Western Africa. This rather inconspicuous species was around for a long time, with the last evidence from rocks showing it lived until 1.65 million years ago. It disappeared from the African continent, but not without leaving a legacy: the Southern Mammoth.

Not a huge mammoth, but still a mammoth. The African Mammoth (Mammuthus africanavus). (Image by Twilight Beasts)

An enigmatic Mammoth
Before we meet the mighty Southern Mammoth, there is an enigmatic mammoth that was the first species to have left Africa (M. rumanus). Fossils have been found in Britain and in sites in Europe, but with only fossil teeth being discovered so far, it remains poorly known. Sadly, the further back in time we go, the fewer fossils we find (land animals do not often preserve as fossils: carcasses are eaten, bones dragged away from the body, bones trampled on). It may have its origins in Africa, around 3.5 million years ago, likely from the First Mammoth, M. subplanifrons. It did manage to make it out of Africa and into Europe, and may have left descendants in the form of the world’s smallest mammoth.

Still hairless, the Enigmatic Mammoth (Mammuthus rumanus) was the first mammoth species out of Africa. (Image by Twilight Beasts)

The Cretan Dwarf Mammoth
On the hot island of Crete, in the early 1900s, the young palaeontologist and explorer, Dorothea Bate discovered some fantastical creatures. An odd sheep that could have been a goat (or was that the other way round?), pygmy hippos, and dwarf mammoths. Yes. A huge oxymoron. But a real creature that once lived. A mammoth that was just 1 meter tall: the Cretan Dwarf Mammoth (M. creticus). How animals get relatively larger or smaller on an island is explained by the Island Rule (or Fosters Rule). Smaller animals on islands are more likely to survive on fewer resources than bigger ones, so natural selection will favour the smaller individuals, and each generation becomes smaller and smaller. Island dwarfism is a fairly common phenomenon and has been seen with dwarf elephants, hippos, deer, and even a species in our own genus, Homo floresiensis.

For a long time, this little animal was thought to have been a mini elephant, descended from the Straight Tusked Elephant. Recent reanalysis by Trowelblazer extraordinaire, Tori Herridge and colleague Adrian Lister, examined in great detail the teeth and showed that it was actually a mammoth! Evidence of its ancestors is sketchy, but with the oldest fossils dating somewhere around 3.5 million years, it is likely to have been the mysterious mammoth, M. rumanus. This was the smallest mammoth to have ever lived. For such a remarkable creature relatively little is actually known about it. The teeth indicate that it ate low bushes instead of grass or tree. With the warm Mediterranean climate, this mini-mammoth was not a hairy beast. It is not known for certain when it became extinct, but there is nothing to suggest that humans were responsible. Despite size, this remarkable little animal was a true mammoth. Small but mammoth!

The smallest mammoth that has ever lived, the mighty-mini Cretan Dwarf Mammoth (Image by Twilight Beasts)

So far that is four species of mammoth: the first mammoth M. subplanifrons, the African Mammoth M. africanavus, the enigmatic M. rumanus and the wonderful mini-mammoth, M. creticus. I wouldn’t be disappointed if that was the end of the line of mammoths. These early species were amazing! Mysterious, huge, tiny: successful animals that once lived and breathed.

Things get even more interesting. Evolution happens a lot faster (relatively speaking) when there are more extreme changes in the environment. Think of when the non-avian dinosaurs disappeared: mammals, birds (avian dinosaurs), and a whole range of other groups radiated into lots of new species in a relatively short time of just a few million years. Animals either cannot cope and die, or are lucky enough to have a random mutation that turns out to be useful. Pure luck can either be the saviour, or demise, of a species. The Pleistocene was full of big changes. This whole Epoch, from around 2.6 million years ago to 10,000 years ago was the Ice Age. It was full of extreme changes in the environment, from periods warm enough to allow hippopotamus to bathe in English Rivers, to periods cold enough for massive glaciers to cover all of Canada. The enormously erratic climate of the Pleistocene was one of the main factors in the evolution of the impressive and iconic Ice Age Beasts. This was no doubt the key driving force for the rapid evolution of the later mammoth species.

The spectacular species of mammoths. Not that there were some early evolutionary dead ends. At many times there were more than one species of mammoth lolloping on Earth. Click on the image to see more detail. (Image by Twilight Beasts)

The Southern Mammoth
And it begins sometime around 2.5 million years ago when one population of the African Mammoth evolved into a wandering giant, the Southern Mammoth (M. meridionalis). With earlier mammoths having a smaller number of ridges on their teeth, the Southern Mammoth had between 12 and 14 ridges on top of low crowns – teeth made for pulverising leaves and shrubs. (Think of a pestle and mortar – the massive teeth of mammoths ground down the vegetation into a slush.) Fossils found with Southern Mammoth remains indicate an environment much different from one you would imagine a mammoth to be at home in: deciduous woodlands. Some specimens at sites in much drier climates (such as Israel and Georgia) have been found with evidence of more grassy environments (similar to the African savannah today). Living in these warmer climates also suggests this giant was not a hairy giant, but likely had similar thick skin to elephants around today.

A flatter higher skull than it’s ancestors and more ridges on those thick molars, the mammoth Southern Mammoth (Mammuthus meridionalis) was a magnificent beast. (Image Twilight Beasts)

The Southern Mammoth was not only capable of adapting to different habitats due to those big, ridged molars, it also travelled very far. Remains have been found in Africa, Europe, Russia and China. This very successful species was on Earth for over a million years. With major changes in the Earth’s climate around 1.5 million years ago, many populations died out. Smaller numbers lumbered along, eventually disappearing over the horizon forever 600,000 years ago.

Before their extinction, one population became standard from other populations. Cold, alone, isolated, this group evolved into a mammoth that would cross an ocean.

The Steppe Mammoth
In the cold, dry grasslands of Siberia, a group of the mighty Southern Mammoth adapted to the cold environment. Around 700,000 years ago we see the first fossils of the enormous Steppe Mammoth (M. trogontherii) descended from this Siberian Southern Mammoth population. With more enamel ridges on those massive molars than its predecessor (up to 19 ridges), the Steppe Mammoth was well suited to eating tougher vegetation on the harsh tundra. Very likely to be the first truly hairy mammoth, this species spread across what was becoming a rich fertile ground for grasses and shrubs and a whole variety of different Pleistocene fauna. They moved West, back over Europe, and even into England, with the famous West Runton Mammoth found in 1990. They also moved east, across Siberia, across the Bering Strait, into Canada, and down into North America.

This was the mammoth that conquered the northern hemisphere.

The largest mammoth of them all, the mighty Steppe Mammoth (Mammuthus trogontherii). (Image by Twilight Beasts)

Sardinian Dwarf Mammoth
The Steppe Mammoth had a huge range across Europe. Some even made it (probably swam) to the Mediterranean island of Sardinia, with later generations becoming smaller and smaller. Around 450,000 years ago, what was once the largest of the mammoth species, quickly evolved into one of the smallest; the Sardinian Mammoth (M. lamarmorai). Isolated from other populations of Steppe Mammoths, the Island Rule forced this mammoth to shrink to a loveable size. It also allowed other wonderful creatures to evolve, including dwarf hippos and giant hares.

Some islands in the Mediterranean were home to dwarf elephants, descended from the large Straight Tusked Elephant, which may be why the Sardinian Mammoth stayed in Sardinia. (If the Sardinian Mammoth managed to make it to another island, there would have been well established populations of dwarf elephants, putting a lot of competition to this newcomer.) This little Sardinian Dwarf Mammoth disappears around 40,000 years ago. Human remains dated to around 13,000 years old have been found on this island, as well as flints that may date much earlier. The extinction of this wonderfully small mammoth may have been due to the arrivals of humans, although without more detailed work on this species, at the moment we don’t know why it vanished for sure.

If you were to call a mammoth cute, the Sardinian Dwarf Mammoth (Mammuthus lamarmorai) would be that mammoth. (Image by Twilight Beasts)

The Woolly Mammoth
Elsewhere in Europe, as the climate shifted once more, the Steppe Mammoth slowly disappeared. Fossil indicate that 400,000 years ago in Siberia it adapted to the changing, bitter landscape, evolving into the most familiar of all the mammoths, the iconic Woolly Mammoth (M. primigenius). These were true ice age beasts. Thick shaggy hair and layers of fat helped protect them from the icy temperatures. And it was cold. An enormous belt of tundra covered northern Europe all the way across to northern America. Called the Mammoth Steppe, this environment was cold but rich in grasses and shrubs to sustain not only Woolly Mammoths, but giant deer, reindeer, woolly rhinoceros and deadly predators like cave lions and sabre tooth cats.

The Woolly Mammoth was at home on this Steppe, a rich ecosystem on the edge of enormous glaciers. With water locked into the great glaciers during the glacial periods, sea levels were much lower, exposing land and connecting continents, and fossils have been found all over the northern hemisphere. Across Siberia the Woolly Mammoths moved down through Canada and into North America. Yep. The Woolly Mammoth was not just a European species.

At least two human species, Homo sapiens and H. neanderthalensis relied on the Woolly Mammoth for survival. Cut marks on bones, burnt bones, cave art, and even evidence of huts made from mammoth bones show that these hairy giants were key to the survival of these humans. All across the northern hemisphere, mammoth populations plummeted towards the end of the Pleistocene. Human hunting had an effect and sadly this coincided with a warming climate. The end of the Pleistocene saw the great glaciers melt, and the Steppe environment slowly retreat north. The Woolly Mammoth clung on to this shrinking environment for as long as they could. One small population managed to survive a little longer on a small island north of Siberia, Wrangel Island. Isolated from predators, this island was their Eden. Until humans arrived. And just 4000 years ago the last of the amazing line of mammoths vanished forever. 4000 years. There were mammoths alive when humans had started writing 5500 years ago. Mammoths were even around when the great pyramids of Egypt were built. They were an incredible group of animals. And they so very nearly made it to today.

The shaggiest mammoth of them all, the one most adored, the Woolly Mammoth (Mammuthus primigenius) (Image by Twilight Beasts)

The Columbian Mammoth
And what of our giant we met at the beginning of this post? Although Woolly Mammoths did make it across to North America, this giant was a different species: it was the enormous Colombian Mammoth (M. columbi). For a long time it was assumed that the enormous Columbian Mammoth evolved from the Southern Mammoth sometime between 1.8 and 1.5 million years ago. A fairly recent relook at hundreds of mammoth teeth in American museums, dispelled this long held theory. The teeth analysed show that they are incredibly similar to the Steppe Mammoth. Lolloping in herds across the cold, grassy tundra, the Steppe Mammoth reached North America, leaving a descendent, the Columbian Mammoth.

The Columbian Mammoth has thrown up more bizarre facts, questioning what we rely on as identifying a species. Palaeontologists look at the remains of animals and plants to work out the diversity of life on the planet. Most of the time these remains are just tiny parts of the whole creature. Specimens of Columbian Mammoth teeth have been analysed using their DNA which strongly suggest that this species was successfully mating with the Woolly Mammoth, leaving fertile offspring. Normally the definition of a species is a group of animals that can breed and produce fertile offspring. (A donkey and a horse can mate, but the mule is sterile.). As we have more data on more recent animals, the line of what we call a species becomes more blurred. We have seen this with recent genetic studies on Neanderthal remains showing that all non-African humans alive today have some Neanderthal DNA.

Sharing its home with other large Ice Age beasts such as giant sloths and sabre-tooth cats, the Columbian Mammoth was one of America’s true giants. After the arrival of humans to America 15,000 years ago, we see the beginning of the end of this beautiful creature. Several sites have recovered human flint implements alongside Columbian Mammoth remains, and bones showing butchery marks. Sadly, a similar story emerges with the Columbian Mammoth sustaining humans which ended the species.

The mammoth in Los Angeles (and most of North America), the Columbian Mammoth (Mammuthus columbi) (Image by Twilight Beasts)

The Pygmy Mammoth
There is one last species of mammoth. In 1856 off the coast of California, about 15 miles out, the small Channel Islands, fossils of very small mammoths were found. This was the Pygmy Mammoth (M. exilis), dating to 80,000 years ago (the third dwarf mammoth species!). Sea levels were a little lower back then, but there was still a vast gulf between the mainland and the islands: a gulf of about 12 miles. Some Columbian Mammoths actually made the journey by swimming! Probably not just on the one occasion, but a few times. Why they decided to take the plunge will never be known. But we do know that they did. Once there, the Island Rule we have seen twice already went into action. Each generation grew a little smaller.

These were not the smallest of the dwarf mammoths, but they were perhaps the most adaptive. Tiny pollen fossils in sediments and the Pygmy Mammoth’s dung show that these little beasts were not limited to one environment on the islands. The youngest fossils date to the same time as evidence for humans arriving, 13, 000 years ago. Isolated for over 60,000 years with no natural predators, the Pygmy Mammoth was vulnerable to human predation, and they too vanished into the twilight.

The third mini-mammoth, not the smallest, but still pretty darn cute. The Pygmy Mammoth (Mammuthus exilis). (Image by Twilight Beasts)

Mammoths have played a part in the lives of humans for tens of thousands of years. Cave paintings in France and Spain beautifully illustrate these creatures drawn from life. Their flesh and bones helped with our own survival. Their bones have also been used as the canvas for some of the earliest pieces of art. Within a breath of the present day the mighty mammoth has vanished forever.

Today there is a lot of attention being given to resurrecting the Woolly Mammoth. Frozen carcasses from Siberia, Asian elephant surrogates, and recreating a Pleistocene Park are splashed across the media every few months. Should we or shouldn’t we is a debate for another post. The mammoths were an incredible group and to see a living giant trundling across the barren tundra landscape, their long shaggy hair floating up with the soft gust of wind, would be a true sight to behold. Personally, we should spend our efforts on protecting the species that are with us today. The last line of an incredible Order of animals are threatened by poaching and habitat loss. If we are not careful, the beautiful elephants too will become another iconic animal our grandchildren will never see, only dream about. The mammoths had their time. Our planet is for the creatures that share it with us today.

Written by Jan Freedman (@JanFreedman)

Our posts on the Proboscidea:

The Steppe Mammoth: The first mammoth of the steppes.

Columbian Mammoth: The last trumpet of a giant.

Woolly Mammoths: Mammoths!

Cloning mammoths: Buttercup the mammoth.

Deinotherium: The Land of the Giants.

Gompothere: The lolloping shovel mouth.

Mastodon: The Nipple Tooth.

Straight Tusked Elephant: An elephant shakes a tree.

Pygmy elephants (Stegodon): Of dwarfs and dragons.

Zyglophodon: Meet Long Tusk.

Further Reading:

Agenbroad, L. D. (2003) ‘New absolute dates and comparisons for California’s Mammuthus exilis.Deinsea. 9(3). pp.1-16.

Agenbroad, L. D., et al. (2007) ‘Mammoths and Humans as Late Pleistocene Contemporaries on Santa Rosa Island.’ Proceedings of the American Geophysical Union. Spring meeting 2007. [Full article]

Agenbroad, L. D. (2010) ‘Mammuthus exilis from the Californian Channel Islands: Height, Mass and Geological Age.’ Proceedings of the 7th Californian Islands Symposium. p.17. [Full article]

Agenbroad, L. D. (2012), ‘Giants and pygmies: Mammoths of Santa Rosa Island, California (USA)’, Quaternary International 255: p.2. [Abstract only]

Bate, D. M. A. (1907). ‘On elephant remains from Crete with description of Elephas creticus sp. n.’ Proceedings of the Zoological Society of London. pp.238-250.

Chang, D., et al. (2017). ‘The evolutionary and phyogeographic histroy of wolly mammoths: a comprehensive mitogenomic anaylisis.’ Scientific Reports. 7:44585. pp.1-10. [Full article]

Ferretti, M. P. (2003), ‘Structure and evolution of mammoth molar enamel’,  Acta Palaeontologica Polonica. 3 48 pp.383–396. [Full article]

Gold, D. et al.. (2014). ‘Attempted DNA extraction from a Rancho La Brea Columbian mammoth (Mammuthus columbi): Prospects for ancient DNA from asphalt deposits, Ecology and Evolution 4 (4). pp.329–336. [Abstract only]

Gunthrie, R. D. (2004), ‘Radiocarbon evidence of mid-Holocene mammoths stranded on an Alaskan Bering Sea island’, Nature. 429. (6993). 746-9. [Abstract only]

Herridge, V. L. & Lister, A. M. (2012) ‘Extreme insular dwarfism evolved in a mammoth.’ Proceedings of the Royal Society Biological Sciences. 279. pp.3193-3200. [Full article]

Lister, A. M. (2004), ‘The Impact of Quaternary ice Ages on Mammalian Evolution.’ Philosophical Transactions: Biological Sciences. 359 (1442). pp.221-241. [Abstract only]

Lister, A, & Bahn, P. (2007), ‘Mammoths – Giants of the Ice Age’, (3rd Edition). London: Frances Lincoln. [Book]

Lister, A. M. & Sher, A. V. (2015). ‘Evolution and dispersal of mammoths across the Northern Hemisphere.’ Mammalian Evolution. 350 (6262). pp.805-809. [Full article]

Lister, A. M., et al. (2005). ‘The pattern and process of mammoth evolution in Eurasia.’ Quaternary International. 126-128. pp.49-64. [Full article]

Lucas, S. G. et al. (1999). ‘Co-occurrence of the proboscideans Cuvieronius, Stegomastodon, and Mammuthus in the lower Pleistocene of southern New Mexico’, Journal of Vertebrate Paleontology 19 (3). pp.595–597. [Abstract only]

Maglio, V. J. (1973), ‘Origin and evolution of the elephantidae.’ Transactions of the American Philosophical Society. 633. pp.1-149. [Full article]

Markoca, A. K. et al. (2013), ‘New data on changes in the European distribution of the mammoth and the woolly rhinoceros during the second half of the Late Pleistocene and the early Holocene’, Quaternary International. 292. 4-14. [Full article]

Martin, P. S. (1999), ‘Twilight of the Mammoths: Ice Age extinctions and the rewilding of America’, university of California Press. [Book]

McDaniel, G. E. & Jefferson, G. T. (2006), ‘Dental variation in the molars of Mammuthus columbi var. M. Imperator (Proboscidea, Elephantidae) from a Mathis gravel quarry, southern Texas’,. Quaternary International. 142-143: 166–177. [Abstract only]

Muhs, D, et al. (2015), ‘Late Quaternary sea-level history and the antiquity of mammoths (Mammuthus exilis and Mammuthus columbi), Channel Islands National Park, California, USA’, Quaternary Research. 83. pp.502-521. [Abstract only]

Patterson, D. B. Mead, A. J. & Bahn, R. A. (2012), ‘New skeletal remains of Mammuthus columbi from Glynn County, Georgia with notes on their historical and paleoecological significance’, Southeastern Naturalist 11 (2). pp.163–172. [Abstract only]

Palombo, M. R., et al. (2012) ‘A reappraisal of the dwarfed mammoth Mammuthus lamarmorai (Major 1883) from Gonnesa (south-western Sardinia Italy).’ Quaternary International. 255. pp.158-170. [Abstract only]

Purdy, B. A. et al. (2011), ‘Earliest art in the Americas: Incised image of a proboscidean on a mineralized extinct animal bone from Vero Beach, Florida’, Journal of Archaeological Science 38 (11). p.2908. [Abstract only]

Shoshani, J. & Tassy, P. (2005). ‘Advances in proboscidean taxonomy & classification, anatomy & physiology, and ecology & behavior’, Quaternary International. 5. pp.126–128. [Abstract only]

Shashoni, J, & Tassy, P. (Eds) (1996), ‘The Proboscidea – evolution and palaeontology of elephants and their relatives.’ United Kingdom: Oxford University Press. [Book]

Stone, R. (2002), ‘Mammoth. The Resurrection of an Ice Age Giant’, Fourth Estate, London. [Book]

Stuart, A. J. (2015), ‘Late Quaternary megafaunal extinctions on the continents: A short review’, Geological Journal 50 (3). pp.338–363. [Abstract only]

Stuart, A. J. et al. (2002), ‘The latest woolly mammoths (Mammuthus primigenius Blumenbach) in Europe and Asia: a review of the current evidence’, Quaternary Science Reviews. 21. 1559-69. [Full article]

Stuart, A. (2005), ‘The extinction of woolly mammoth (Mammuthus primigenius) and straight-tusked elephant (Palaeoloxodon antiquus) in Europe’, Quaternary International. 126. 171-7. [Abstract only]

Sutcliffe, A. J. (1985), ‘On the track of Ice Age mammals‘, British Museum (Natural History). [Book]

Tichonov, A, & Burlakov, Y. (2008), ‘Causes of Northern Giants’ extinction.’ Science in Russia. (Moscow: Nawka). 2. pp.48-53. [Abstract only]

Wei, G. B., et al. (2010), ‘New materials of the Steppe Mammoth, Mammuthus trogontherii, with discussion on the origin and evolutionary patterns of mammoths.’ Science China Earth Sciences. 53(7). pp.956-963. [Full article]

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The Stuff Of Night-Mares

Now that my thesis is nearly at the end, I can tell you all this secret. I have been so tiddled off at the archaeological paradigm that all ancient horses were wee tiny ponies. Victorian zoologists like Ridgeway and Ewart made up stuff about types, which certainly were not formalised breeds, linking everything they liked, and very much approved of, to the exquisite Arabian horse, and sometimes they made up a whole new species for themselves (like Equus caballus celticus – a totally tiny Celtic pony!) because it seemed like a good idea at the time. However, they were two smart cookies for their time, so I will forgive them, as any horsie person knows, horses get under your skin somewhat, and if we can imagine unicorns, we can imagine special Celtic ponies!

However, way out east in the Iron Age, horses were horses, with the ‘fossil’ breed of the Akhal Teke, and the bigger cold bloods, as they call them in equestrian circles. Now, don’t get me wrong – there were indeed very little ponies in the past. Insular breeds, such as the British Exmoor and Scottish Eriskay, tend to be small, and humans across Europe and Eurasia bred the wildness out of the true wild horse, the Tarpan, making the overall size of equids smaller and narrower and presumably easier to control. The now-extinct Lofoten island pony was likely the ancestor of the real Shetland pony, the ones which haven’t been bred up with new bloodlines chucked in every now and then. The Lofoten was a very small, hairy pony breed, like a pure white teddy bear, which came to a rather instant extinction, when the very last one was shot deliberately in 1897 to preserve its body for science. I know, right?  The poor little thing has been on display since then, in Bergen Museum.


The last of his line – the Lofoten pony, which was made extinct to be a museum piece – Victorians, eh! ( image from Ridgeway 1905)

Nature, however, loves diversity – life forms adapt and grow, or shrink depending on a whole host of environmental factors. If you went back to the Pleistocene, you’d find lots of wild equids of all sizes and all shapes….Equus alaskae, E. nevadanus, E. taeniopus… the list is massive. But among them, there was a giant, possibly bigger than a Clydesdale –  Equus giganteus. There are some pretty large gaps of knowledge about E. giganteus, because most of the evidence is based on just teeth. There was supposedly also a full mandible or skull found by ‘Mister Bones’, the 19th century fossil hunter Barnum Brown.

The early days of fossil collecting was often met with a flurry of new species. With just a few fragments to go on, later palaeontologists have since noticed that many fossils of ‘new species’ actually all belong to just one species. There’s confusion over whether the fossils of  E. pacificus might be a variation of E. giganteus, and there might even be a larger horse,  E. enormis, although there is even less evidence for it, being confined to some West Coast USA fossils detailed in Murray’s 2008 thesis. They’ve been primarily found where there were Pleistocene prairies – Texas, Kansas, Oklahoma, California and Nevada. Imagine how stunning it would have been to watch herds of these creatures grazing and galloping. Mind you, with their considerable size, they would need a fair bit of space to gallop wild and free.

Now, we measure horses at the withers – the little lump at the base of the neck, not the tops of their heads and we call it hand high, usually abbreviated to hh. E. giganteus likely measured over 20 hands, or 2m (6’7”), as simply, we don’t even know if the few scant remains of teeth we have are from juvenile specimens. (Estimates on total height are 10 feet!) However, we know enough to speculate on  E. giganteus to make it a creature of a Tolkeinesque dreamtime, bigger than a Shire Horse, and weighing in between 700 and 1600 kg depending on which reconstruction model you choose to go with (palaeontology is, in this respect, every bit as bad as archaeology- everything has a ‘but what if’ added!).  Much good, solid horse sense was written on this by Gidley in 1901, who stated that the superior molar tooth found in Texas in the 19th century was bigger by over a third of the diameter than that of an equivalent modern draft horse.

The ginormous size of the North America giant horse, Equus giganteus.

While we can pretty much reconstruct the big, dignified head of E. pacificus, another giant Pleistocene horse of the Wild, Wild West, we cannot yet do so for E. giganteus. But we can make some pretty good guesses. So-called ‘primitive’ types and breeds usually have certain distinctive markings, fingerprints from the deep past, carried on shoulders and soft muzzles to remind us of their ancestors, who existed before humans domesticated them. Have a look at the Przewalski horse, with its big solid head, so like cave drawings of Chauvet, or the Dulmen from Germany, or the wee English Exmoor. They have what we call a ‘mealy muzzle’, in that their soft noses are often paler than their coat colouring while their hocks, or lower legs , are darker. Across their withers, running down their backs, is a dark line of hair called a dorsal stripe, although I think of it as a time line, proudly stating the ancient ancestry of the breed. On occasions, with very old breeds, you may even get a hint of stripes on the lower limbs. I’d suspect that Equus giganteus had all of the above, loud and proud. They do say everything is bigger in Texas!

Our giant horses were present in the great Pleistocene grasslands of the western USA, but probably took their first leggy steps in the Pliocene, around 2.5 million years ago. Horses are believed to have evolved in the North American continent, moving into South America about 1.5 million years ago. All American species of wild equids became extinct by 10,000 years ago, and the prairies and grasslands were empty of neighs and thundering hooves until the reintroduction of Equus caballus by the Spanish conquistadores of the 1500’s. Some of their horses escaped and reclaimed the land as mustangs, the iconic feral mixed-breed types. But they were, and are, normal-sized horses, unlike the big beauty of E. giganteus.


A friend of mine, Maximus, a fjord pony, who happens to be showing off his  ancient ancestry rather well, with his lovely mealy muzzle and hog mane (he’d like to be your friend too if you head up Armoy direction)

So, what brought the reign of these giant, wild spirits of the grasslands to an end? And when exactly did it happen? The truth is, we really don’t know yet. Regular readers will know that there were huge climate changes occurring during the Pleistocene, which were responsible for some of the megafaunal extinctions. The climate shift resulted in vegetation changes, from higher protein plants ( known as C3 vegetation) to coarser, harder to digest grasses ( known as C4 grasses) resulted in the grasslands being incapable of sustaining many megafaunal species.This is usually considered the reason for  extinctions of smaller horses of the Americas, and it may be that E giganteus  left the land earlier. After all, there was a whole lot of horse to feed.

However, the time we estimate the great horses vanished coincides with the arrival of the Clovis people in North America, probably around 13,000 years ago. Clovis points, the lethally sharp lithics used by these ancient people have been found in Texas although, so it is anyone’s guess as to whether the horses were hunted into extinction. We know humans hunted horses in Europe, so while there is no definite early  evidence yet, it wouldn’t be so surprising if a future excavation in the southwest states turns up something more conclusive than ‘maybe’.  An indirect possibility is that fires caused by humans could well have destroyed much of the original grasslands. Remember, with large numbers of herbivores on such grasslands, you tend to get predators too, and they will be quite happy to pick off an unwary human as much as any other smaller creature on the prairie.Short-sighted solutions of slash and burn may have hastened the end of our lovely horse.

So, we have no idea when Equus giganteus became extinct. Recently, in the name of research, I had the chance to gallop out on a Clydesdale, which is still smaller than E. giganteus. Her name was Niňa, which, meaning little girl, was a bit of a joke. She was huge. I’m an experienced horsewoman, but galloping her felt as if I were strapped to the outer engine of an aircraft taking off, like straddling thunder.  While all wild horses are enchanting, triggering something deep inside us as humans, I can’t help thinking of  what perhaps dozens or hundreds of horses, well over 20 hh, would be like – storm-clouds made flesh, whinnying and snorting across the grasslands. What I’d give for a TARDIS ( and probably a saddle and bridle) !

Written by Rena Maguire (@JustRena)

Further reading:

Baskin, J. A., 1991. ‘Early Pliocene horses from late Pleistocene fluvial deposits, Gulf Coastal Plain, South Texas’.  Journal of Paleontology. 65. pp. 995-1006. [Abstract only]

Cantalapiedra, J.L., Prado, J.L., Fernández, M.H. and Alberdi, M.T., 2017. Decoupled ecomorphological evolution and diversification in Neogene-Quaternary horses. Science355(6325), pp.627-630. [Full article]

Cope, E. D., 1891. ‘On a skull of Equus excelsus Leidy, from the Equus beds of Texas’ American Naturalist. 25. pp 912-913.

Cope, E. D., 1885 ‘Pliocene horses of southwestern Texas’  American Naturalist. 19. pp1208-1209.

Dalquest, W.W., 1977. ‘Mammals of the Holloman local fauna, Pleistocene of Oklahoma’. The Southwestern Naturalist, pp.255-268. [Full article]

Dingus, L and Norel. M 2010. Barnum Brown; the man who discovered Tyrannosaurus Rex. Berkley: University of California Press

Eisenman, V. n.d Giant Horses. (Full article)

Gidley. J.  1901. Tooth Characters and Revision of the North American Species of the Genus Equus. Bulletin of the American Museum of Natural History 14(9):1-60. [Full article]

Grayson, D.K. and Meltzer, D.J. 2003. ‘A requiem for North American overkill’. Journal of Archaeological Science30(5), pp.585-593. [Full article]

Haynes, G., 2014. ‘North American Megafauna Extinction: Climate or Overhunting?’. In Encyclopedia of Global Archaeology  New York: Springer. pp. 5382-5390. [Abstract only]

Hill ME, Hill MG and Widga CC. 2008. L’ate Quaternary bison diminution on the Great Plains of North America: Evaluating the role of human hunting versus climate change’. Quaternary Science Reviews 27(17-18). pp 1752-1771. [Full article]

Lambert WD and Holling CS. 1998. ‘Causes of ecosystem transformation at the end of the Pleistocene: evidence from mammal body-mass distributions’. Ecosystems 1(2) pp 157-175. [Abstract only]

Lyons RK and Hanselka CW ‘Grazing and browsing: how plants are affected’. Texas Cooperative Extension. Texas A&M University System. [Full article]

Murray, L. K. 2008. Effects of taxonomic and locality inaccuracies on biostratigraphy and biochronology of the Hueso and Tapiado formations in the Vallecito Creek–Fish Creek Section, Anza-Borrego Desert, California. PhD Dissertation, University of Texas at Austin, pp. i-xxiii, 1-532. [Full dissertation]

Owensby CE. 1998. ‘Role of grasslands as modifiers of global climate change’. Proc. 18th International Grassland Congress. pp 9-12 [Full article]

Ridgeway, W. 1905. The Origin and Influence of the Thoroughbred Horse. Lofoten pony reference  pp 119-121.  [Full article]

Stromberg MR, D’Antonio CM, Young TP, Wirka J, Kephart PR. 2007. ‘California grassland restoration’. in: Stromberg MR, Corbin JD and D’Antonio CM(eds) California grasslands: Ecology and management. Berkeley, Los Angeles, London: University of California Press. pp 254-280. [Book]

Posted in Horse | Tagged , , , , , , , , , , , , , , , | 4 Comments

The Devil on the mountain

When hiking high in the Alps you may encounter the devil himself. Or so claims Swiss naturalist and mountaineer Horace-Bénédict de Saussure (1740-1799), when he writes, that one will be only ‘in the company of the devil,…’ if climbing the Alpine peaks. De Saussure was  indeed describing a horned animal with brown to black fur, but despite its appearance, the chamois is not really a demonic entity. The chamois is a gracile, small ungulate, native to many European mountains, with characteristic backwards hooked horns. Like real goats, chamois are excellent climbers. The flexible, cloven hooves are well adapted to move on rocky and rugged terrain. With a soft inner hoof-pad and a hard outer rim, the hooves will find grip even on the steepest cliffs. The name chamois derives from the Greek kemas, a name given in antiquity to wild goats. Kemas itself derives from kamp, an ancient Indian word for jumping. Seeing a chamois jump from rock to rock one can understand why this name fits it so well. However, some anatomical peculiarities separate this animal from real goats, for example, male chamois have no beard. The famous Gamsbart, the ‘chamois-beard’ displayed by hunters on their traditional hat, is made from a tuft of long hairs, growing on the back and rear end of male chamois during the rutting season. Early zoologists described the chamois as a European antelope species. However, today the chamois is classified as mountain-goat in an own genus, named Rupicapra.

A chamois in its natural habitat. (Photo by D. Bressan)

The chamois plays an important role in Alpine folklore. In medieval times the Alps were often simply referred as Gamsgebirg, the chamois mountains, and still today many names of mountains or peaks refer to this animal. Some legends tell that the chamois was indeed created by the devil, to lure young hunters to their death in the mountains. Other times, so says a legend from the Italian Dolomites, the devil himself, disguised as a pitch-black chamois, appeared to the terrified hunters.

The hunt for chamois was indeed very dangerous. Hunters used ropes to catch the animals or long spears to push them from steep cliffs. Many stories tell about hunters who fell to their death. With the invention of rifles, the hunt became much easier and safer. Between 1700-1850 the species became rare in the Alps. De Saussure writes in 1780 ‘Though the profit is small, the people of Chamouni hunt … with passion, and so these creatures are diminishing in number in the most noticeable fashion. … The hunters of Chamouni have already utterly destroyed or driven away the bouquetins [ibex], which, once were common on their mountains, and it is likely that in less than a century neither chamois nor marmots will be seen.’ In the 19th century, the chamois was locally extinct, but fortunately, with the introduction of modern hunting rules and the creation of protected areas, nowadays the chamois is again common in the Alps.

Finnish vertebrate paleontologist Björn Kurtén (1924-1988) referred to the origin of the genus Rupicapra as a mystery. Today chamois populations are found in the European Alps, Apennines, Pyrenees, Carpathian Mountains and the French Massif Central (and introduced in New Zealand), even if it the genus is not considered of European origin. Fossils are indeed very rare and the chamois seems to appear suddenly in the fossil record during the last interglacial. Probably the ancestors of the three modern chamois species, the Alpine-Chamois R. rupicapra, the Pyrenean-Chamois R. pyrenaica and the Apenninic-Chamois R. (pyrenaica) ornata, migrated during the Riss glaciation (250,000-150,000 years ago) from Asia into Europe, becoming abundant here during the last ice age (80,000-12,000 years ago) and colonizing all mountain ranges in central Europe.

Living in mountains helps the chamois to evade most predators, with the exception of humans and eagles. The fossilization of bones is difficult in such a terrain because carcasses are exposed and scavenged. Fossils are therefore known only from bone accumulations in caves. As are fossils rare, so is the prehistoric art showing the chamois. Maybe this animal, living on inaccessible terrain, was a difficult and not very appreciated trophy for prehistoric hunters. There was just no interest to depict it. The rare examples we have are however quite intriguing. There are only six sites of cave art known to date, all located in France or Spain. One example is a single chamois, shown in a painting of red ochre in the Cave of La Pasiega, Spain.

The chamois in the Cave of La Pasiega, image from H. Obermaier, ‘Der Mensch der Vorzeit’ (1912)

Quite more common are depictions in prehistoric small, ‘mobile’ artwork. An entire chamois herd was found as engraving on a reindeer-bone. Discovered in the cave of Gourdan, France, the scene maybe shows a lion, spotted in the background, chasing the herd (modern chamois are still highly gregarious, forming large herds during the winter).

The chamois of the Cave of Gourdan, image from H. Osborn, ‘Men of the Old Stone Age’ (1916)

In one curious example found in the Mas d’Azil Cave, a young chamois (or maybe an ibex), carved into a spear-thrower made from horn, looks back at birds, picking on its own feces. Another mysterious discovery involves a small bone-disc with engravings on both sides, discovered in the archaeological site of Laugerie-Basse. Believed to be an early example of a bottom, its exact use is unknown. It was suggested that it was used as decoration on spindles, used for spinning yarns from natural fibers. One side shows a standing animal, on the other side, the animal seems to lie on the ground. If the disc is spun quickly enough, the chamois seems to repeatedly stand up and lie down. Maybe this is the oldest known example of an animation known to date, with an age of 12,000 years.

However, the most enigmatic depiction are the ‘chamois dancers’, found as an engraving on a mammoth tooth, discovered in the Abri Mège, Teyat in France. The three little chamois with human legs could be animal spirits or shamans, wearing the animal’s fur in some sort of ritual. It is also possible, that the engraving shows a hunting scene. Like some modern hunters, hunters 17,000 years ago could have approached a herd by disguising themselves as animals.

The ‘chamois dancers’, image from H. Obermaier, ‘Der Mensch der Vorzeit’ (1912)

At the end of the last ice age, when the treeless mammoth steppe was slowly replaced with forests, the chamois, well adapted to cold environments, followed the receding glaciers into the mountains. There the various populations became isolated. The modern spotted distribution and the various recognized species are the results of this isolation and local evolution. However, this tough ice age survivor is nowadays threatened by tourism and climate change. In some areas, way too many tourists have pushed the chamois from its habitual pastures to less suited grounds. A resent study showed that as temperatures in the Alps continue to rise, the average body size of young chamois tend to shrink. It is not entirely clear if this is the direct result of chronic undernourishment, as high temperatures cause stress in the animals, they have to rest more and can dedicate less time searching for food. Diseases and parasites, like the feared mange, caused by parasitic mites, can spread better with warm temperatures.

Despite this, it is still quite easy to spot herds of chamois in the Alps. One should just be aware if encountering a pitch-black chamois. It could be a coal-chamois, as a rare variety of too intense pigmented chamois is called, maybe a shaman in a fur costume, or maybe it really is the devil in disguise…

Chamois costume of a traditional folklore carnival in the Alps. (Image by D. Bressan)

Written by David Bressan (@David_Bressan)

Edited by Jan Freedman (@JanFreedman)

Further Reading:

KURTEN, B. (2007): Pleistocene Mammals of Europe. Aldine De Gruyter Publisher: 317. [Abstract]

LOVARI, S. (1987): Evolutionary aspects of the biology of chamois, Rupicapra spp. (Bovidae, Caprinae). The Biology and Management of Capricornis and Related Mountain Antelopes: 51-61. [Abstract]



Posted in Cave art, Chamois | Tagged , , , , , , , , , , , , , , , , , , , , , | 1 Comment

An elephant shakes a tree

Mammoths get all the attention. Like an annoying younger sibling, they hog a limelight that should be more equally shared. Occasionally, the mastodon gets a whisp of publicity, which is notable mostly for its rarity. The vast family of proboscideans barely get a look in. When did you last hear any news about exciting new finds of Stegodon or Cuvieronius or Sinomastodon or Notiomastodon or Stegomastodon? What about Palaeoloxodon?

I thought so.

Skeleton of Palaeoloxodon antiquus in the Museo paleontologico of the university Sapienzia, Rome. Public Domain Image via Wikimedia Commons.

Reconstruction of Palaeoloxodon antiquus and calf. Image by Apotea via Wikimedia Commons.

Palaeoloxodon antiquus should be better known. This species, also called the straight-tusked elephant, was found all over Eurasia during the warm periods of the Pleistocene. Even in Britain. We’ve got bones of this giant from interglacial gravels underneath Trafalgar square, on the east Anglian coast, and from right under the route now taken by the Channel Tunnel rail link. In Europe, Palaeoloxodon even submarined its way to some of the Mediterranean islands and evolved into unique, teeny-tiny forms on the islands of Malta, Cyprus, Tilos, and Sicily (mammoths did something similar on the island of Crete). In fact, if you close your eyes and squint real hard, there is some controversial evidence that these dwarf elephants may have squeaked into the historical period. A very interesting panel from the tomb of Rekh-Mi-Re, an 18th dynasty vizier, buried in Thebes, shows what appears to be a fully mature elephant with tusks, but only waist high. Could it have come from a Mediterranean population? Possibly, but it could also just be a stylistic convention. We don’t have a good idea of when the insular elephants went extinct but it was most likely sometime during the middle Holocene, with some perhaps hanging on until the Bronze age.

Size comparison of the dwarfed Tilos elephant compared to mainland Palaeoloxodon antiquus. Image by A. Mangione from Masseti 2001

Elephant painting from the tomb of Rekh-Mi-Re. It has also been used to suggest a trading route between ancient Egypt and Wrangel Island(!). Image from Masseti 2001

We do have a better idea of when the fullsized straight-tusker disappeared. It seems to have been widespread during the height of the last interglacial when temperatures were comparatively balmy. It shared the European landscape with hippos (Hippopotamus amphibius), Merck’s rhino (Stephanorhinus kirchbergensis) and other typically warm-adapted fauna. It probably wasn’t as hirsute as its woolly cousin and it’s helpful to picture the living animal as an Asian elephant on steroids. Four metres to the shoulders, longer tusks, just mammoth, mastodonic, huge overall. When the climate started to cool again, it probably retreated to some of the classic Pleistocene refugia: Iberia, Italy, perhaps the Balkans too. Some evocative footprints attributed to P. antiquus have been excavated in Portugal from fossilised sand dunes. However, even these regions weren’t warm enough and the mainland straight-tusked elephant was probably gone by the end of the Middle Pleistocene. Although, having said that, there are radiocarbon dates for late Pleistocene Palaeloxodon in Portugal, but these haven’t convinced everyone. Even more fringe is the idea that straight-tusked elephants were roaming China during the historical period. This is based on analysis of bronze artefacts which show elephants with unusual trunk features. But again this could just by stylistic convention.

An early Chinese bronze depiction of an elephant. It’s claimed that the trunk tip contains two fingers- a feature found in African elephants and mammoths but not Asian elephants. Image from Li et al. 2011

Right from the start, researchers have been pretty clear about where the affinities of P. antquus lay. It started off as Elephas antiquus; Elephas is the genus of the Indian elephant (Elephas maximus). However, as is its wont, ancient DNA has now come along and mixed things up a little. This week Meyer et al. published Palaeoloxodon antiquus mitochondrial genomes and nuclear DNA from Germany and it has turned things on their head. Despite reams of morphological data suggesting the straight-tusker and the Asian elephant were sister species, the DNA data puts Palaeoloxodon squarely with the African elephants. Not only that but it puts it as sister to the African forest elephant, Loxodonta cyclotis. This means that there is more genetic distance between the African savannah elephant (L. africana) and the African forest elephant that are found today, than there is between the forest elephant and Palaeoloxodon antiquus.

Above: Comparison of the phylogenies produced from mitochondrial DNA and nuclear DNA, showing their high support and congruence. Below: Reconstructed elephant family tree, based on the new data. Images from Meyer et al.

This is important. If you’ve paid any attention to the news recently then you know that elephants are in serious trouble. As in, we are killing them all. Poachers and ivory traders don’t give a shit about the difference between savannah and forest elephants, but we should. The destruction of wild elephants may be pushing forest elephants to the brink of extinction yet we can still look at the savannah elephants and think we have time to save them. We don’t. The continued loss of forest elephants represents the loss of a distinct lineage that needs to be recognised as a different species. Hopefully the reshuffling of Palaeoloxodon will help to show that the two African elephants are distinct, unique species and conservation measures can be tailored to their distinct, unique needs.

Elephant killed by poachers. Image by Ina96 via Wikimedia Commons.


Written by Ross Barnett (@DeepFriedDNA)

Further Reading:

Binladen, J., M. T. Gilbert, and E. Willerslev. “800,000 Year Old Mammoth DNA, Modern Elephant DNA or PCR Artefact?”. Biol Lett 3, no. 1 (Feb 22 2007): 55-6; discussion 60-3.[Abstract]

de Carvalho, C. N., S. Figueirido, and J. Belo. “Vertebrate Tracks and Trackways from the Pleistocene Eolianites of Sw Portugal.” Comunicações Geológicas, no. 103 (2016): 101-16.[Full Text]

Franks, J. W. “Interglacial Deposits at Trafalgar Square, London.” The New Phytologist 59 (1960): 145-50.[Abstract]

Herridge, V. L., and A. M. Lister. “Extreme Insular Dwarfism Evolved in a Mammoth.” Proc Biol Sci 279, no. 1741 (Aug 22 2012): 3193-200.[FullText]

Li, J., Y. Hou, Y. Li, and J Zhang. “The Latest Straight-Tusked Elephants (Palaeoloxodon)? “Wild Elephants” Lived 3,000 Years Ago in North China.” Quaternary International 281 (2011): 84-88.[Abstract]

Masseti, M. “Did Endemic Dwarf Elephants Survive on Mediterranean Islands up to Protohistorical Times?” In The World of Elephants – International Congress, 402-06. Rome, 2001.[FullText]

Meyer, M., E. Palkopoulou, S. Baleka, M. Stiller, K. E. H. Penkman, K. W. Alt, Y. Ishida, et al. “Palaeogenomes of Eurasian Straight- Tusked Elephants Challenge the Current View of Elephant Evolution.” eLife, no. 25413 (2017): 1-14.[Full Text]

Orlando, L., M. Pages, S. Calvignac, S. Hughes, and C. Hanni. “Does the 43 Bp Sequence from an 800,000 Year Old Cretan Dwarf Elephantid Really Rewrite the Textbook on Mammoths?”. Biol Lett 3, no. 1 (Feb 22 2007): 57-9; discussion 60-3.[FullText]

Palombo, M. R. “Endemic Elephants of the Mediterranean Islands: Knowledge, Problems and Perspectives.” In The World of Elephants – International Congress. Rome, 2001.[FullText]

Poulakakis, N., A.P. Parmakelis, P. Lymberakis, M. Mylonas, E. Zouras, D. S. Reese, S. Glaberman, and A. Caccone. “Ancient DNA Forces Reconsideration of Evolutionary History of Mediterranean Pygmy Elephantids.” Biology Letters (2006).[Abstract]

Stuart, A. J. “The Extinction of Woolly Mammoth (Mammuthus Primigenius) and Striaght-Tusked Elephant (Palaeoloxodon Antiquus) in Europe.” Quaternary International 126-128 (2005): 171-77.[FullText]





Posted in Columbian Mammoth, Deinotherium, Extinction, Hippopotamus, Mastodon, Palaeoloxodon, Stegodon, Steppe Mammoth, Woolly Mammoth | Tagged , , , , , , , , , , , | 4 Comments

On the shoulders of giants

In 1835 the young, and somewhat cavalier, Charles Darwin landed for the first time on the Galapagos archipelago. As well as sending hundreds of specimens back to England, Darwin enjoyed exploring the islands and watching the local species in their natural habitat. Arguably one of the most magnificent animals he set eyes on were the giant tortoises. In his zoological and geological write up, The Voyage of the Beagle, he dedicates quite some space to these enormous reptiles.

Darwin observed them and studied them closely.

He also rode them.

Yep. Charles Darwin rode on the back of giant tortoises.

“I frequently got on their backs, and then giving a few raps on the hinder part of their shells, they would rise up and walk away;- but I found it very difficult to keep my balance.”* (Darwin, 1839)

Giant tortoises are pretty big. Big enough to straddle. (Please don’t ever straddle one.) On the several islands of the Galapagos archipelago there were 13 different species of giant tortoises, with at least 2 of these only recently becoming extinct. What’s more amazing is that the giant tortoise is not just restricted to the Galapagos archipelago. Another species lives on the Aldabra atoll, in the Indian Ocean: the Aldabra giant tortoise (Aldabrachelys gigantea). Until just a few hundred years ago there were giant tortoises on several islands in the Indian Ocean, before humans arrived.

Not all giant tortoises live in the Galapagos archipelago: the Aldabra giant tortoise (Aldabrachelys gigantea) from the Seychelles.(Image Public Domain)

Once there were many more species slowly lumbering around. And slowly lumber they did: they started to vanish around 100,000 years ago with most becoming extinct by around 10,000 years ago. It seems they were easy pickings for one species of bipedal ape. Our beast was pretty similar to the Galapagos giant tortoise, only, much, much bigger. This was the largest giant tortoise ever: Megalochelys atlas.

This was a very successful genus spanning the Miocene (around 5 million years ago) to the Late Pleistocene (around 10,000 years ago), living in India, Pakistan, Indonesia, and possibly as far west as Europe. With a similar anatomy to the Galapagos giant tortoise, this extinct beast likely ate similar food: dried leaves on the forest floor with the huge stocky limbs holding up their enormous house.

The enormous giant tortoise Megalochelys atlas shows just how big it was compared to Rey. (Image by Jan Freedman)

But why did they get so big? Familiar gigantism in animals is seen on islands (unimaginatively known as island gigantism). Here, isolated on an island a normally small mainland animal becomes pretty darn big. We have seen this in a number of our beasts: the dodo (being an oversized pigeon); the giant lemur of Madagascar; the huge Hawaiian Duck; the massive flightless Moa; and many more. Some animals will quickly get bigger in their new environment due to lack of predators and new and plenty foods around.

These ginormous tortoises didn’t live on small islands; they lived on a continent. Animals can grow pretty big on large landmasses too. You might be familiar with those pesky animals that get all the media love: dinosaurs. Some of these grew to unfathomable sizes for animals, where each new discovery pushes the limits to how big an animal can get. Proboscids, like Mammoths and the giant Zygolophodon, also grew to massive sizes. Perhaps there is no specific reason for these tortoise growing so large, except that they could. In good times species flourish, and Miocene Eurasia was a rich fertile environment which doubtless supported these (and many other) incredible creatures. Today we find the giant tortoises isolated on islands and assume this is how they got their crazy big sizes. But they got big on the mainland first, and later arrived on islands by chance.

Despite first being discovered back in 1844 there is not that much information about this massive giant tortoise. There is no mention of it in any of my Pleistocene books on my bookshelf, and relatively few references in the literature. They were very likely similar to their island relatives in lifestyle. The giant tortoises on the mainland became extinct between 100,000 and 10,000 years ago, likely by the hand of humans although no roasted shells have yet been discovered.

What saved the giant tortoises alive today from extinction were the islands they made home. But even then soon after humans arrived many island populations vanished forever. When Darwin was on the Galapagos, he notes how sailors brought dozens of giant tortoises down to the ship with relative ease. Previous years, they were so abundant that sailors had caught around 600 tortoises. They were so easy to catch because they relied on their size to protect them and they had no reason to fear humans. These giants were a favourite of sailors to take back for fresh meat on their long voyage home: the tortoise lived on the ships without the need of much attention, and gave the crew fresh meat when needed. They were easy pickings for sailors, and no doubt that Megalochelys atlas was easy pickings for the first Homo sapiens who encountered them in Eurasia. I often wonder if any of these people were gutsy enough to hop on and straddle the biggest tortoise to have ever lived.

Written by Jan Freedman (@JanFreedman)

Further Reading:

Arnold, E. N. (1979), ‘Indian Ocean giant tortoises: Their Systematics and Islands Adaptations,’ Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 286: 1011, The Terrestrial Ecology of Aldabra (Jul. 3, 1979), pp. 127-145.

Badam, G. L. (1981) ‘Colossochelys atlas, a giant tortoise from the Upper Siwaliks of North India,’ Bulletin of the Deccan College Research Institute. 40. pp.149 – 153.

Falconer, H. & Cautley, P.T. (1844). ‘Communication on the Colossochelys atlas, a fossil tortoise of enormous size from the Tertiary strata of the Siwalk Hills in the north of India.’ Proceedings of the Zoological Society of London. 12. pp.54–84.

Hansen, D. M., et al. (2010). ‘Ecological history and latent conservation potential: large and giant tortoises as a model for taxon substitutions.’ Ecography. 33:2. pp.271-284.

West, R., Hutchinson, H., & Munthe, J (1991) ‘Miocene vertebrates from the Siwalik Group, Western Nepal,’ Journal of Vertebrate Palaeontology. 11: 1. pp.108-129.

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The Evolution Underground

My daughter froze. She had just been hopping across the coarse, sun-baked sand bank to paddle and splash in the cold, clear River Teign. Giggling, she skipped back across the gravely sand, where she would turn and repeat the process, as she had already done a dozen times to her delight. Only this time she came to an abrupt halt. She had spotted something. Her older brother had taught her well: a little movement or a flash of colour may be some hidden insect just waiting not to be discovered. She crouched down and lay on her tummy. She was transfixed. I lay down beside her and that’s when I saw it: a small, rather dull bee digging in the sand.

She had watched big, hairy bumble bees disappear inside beautifully coloured petals to emerge covered in yellow dusty pollen; a small price to pay for that rich nectar. She had seen bee keepers, looking like some extra in a 60s sci-fi alien film, pulling honey from human made bee hives. She had even seen a chimpanzee easily destroy a bee hive with a big stick on a natural history programme. For a three year old, she knows quite a bit about bees. But she had never seen a digging bee.

A very funky, 80s looking mining bee (Andrena haemorrhoa) (Image by Charles Sharp Public Domain)

This ‘digging bee’ was actually a bee belonging to the largest group of solitary bees: the mining bees. They dig out a little burrow in sandy soil, where they lay around five eggs in individual pods in the sides of the burrow. Mining bees are cool because they actually dig tunnels. They also help pollinate plants along with beetles, flies and plenty of other mini-beasts. Another cool thing is that some species are hosts for other animals. In Britain the beautiful, big, shiny Oil Beetles rely on the mining bee to survive: the youngsters jump onto the back of their favourite type of mining bee and catches a ride back to it’s nest. Once there it devours the eggs, and uses the mining bee’s burrow as a home for safety against predators until it emerges as an adult. What a fascinating dependence for a species to survive: without one specific species of mining bee, this oil beetle wouldn’t be here.

The hidden world of life under our feet is often out of sight, and so out of mind. Bees digging burrows? Ferocious beetle larvae using these burrows as a surrogate home? This unseen world is pretty incredible. It turns out that this world is around us more than we realise. And it has been hidden for a very long time.

I had always had a little fascination for this hidden world where many animals actually spend the majority of their lives, to sleep, mate, and bring up their little ones. I recently saw on Twitter that a fellow science tweeter, Anthony Martin, had a new book just released: The Evolution Underground. Because Twitter is all about networking, collaborating, and sometimes being politely cheeky, I asked if I could review a copy. And because the science community on Twitter is wonderful, I was sent a copy. Although this book is not solely about Pleistocene burrows, I wanted to review it because the story of burrows is so often forgotten about in daily life.

I wanted to find out more about burrows and how there could be an entire book about them.

And this book is all about the burrows. All. About. The. Burrows.

A pretty impressive slab of rock holding huge burrows made by crustaceans, from the Middle Jurassic. (Image by Mark Wilson. Public Domain)

You may think that this might be pretty dry reading, but you will be very surprised. If you hadn’t thought burrows painstakingly dug out by animals were interesting, you haven’t seen a burrow painstakingly dug out by an animal. Martin, now at Emory University, Atlanta, has spent his professional career studying fossil tracks and burrows, so he knows an awful lot about them (this field of palaeontology is called ichnology). This shines through as Martin deftly describes dimensions of shapes, twists and turns of more burrows than I ever imagined existed. These remains of evidence of animals, such as footprints or burrows (known as trace fossils), can tell us an awful lot about life: even life that vanished hundreds of million years ago.

I could happily give a summary of each chapter because they are individually fascinating, all with their own unique little stories. But I will keep some of the suspense and discoveries for you. However, I will tell you about my two favorite chapters looking at burrows: one delving into deep time, and another I am much more familiar with.

Before 540 million years ago, when the land was naked and the seas were bustling with strange and wonderful forms, life was very different. The fossils from just before this time were just plain odd. Collectively known as the Ediacaran Biota, these are some of the most bizarre fossils you will ever see, they don’t really look like an animal or plant: they don’t look like anything really. Bizarre, but also spectacular. These faint imprints are the earliest evidence of multicelluar life on our planet, from between 635 and 544 million years ago. I was delighted to find a chapter dedicated to the Ediacaran time and just after, and more so when I read about the work of a colleague I recently met from across the pond.

One of the most beautiful slabs of rock I have ever held. Ediacaran fossils at the North Carolina Museum of Natural Sciences. On it you can see imprints towards the top left, and more elongated shapes towards the bottom right. Are these burrows or fossils?  (Photograph reproduced with permission from Antony Martin)

I met Trish Weaver some months ago whilst visiting several museums on a mini-museum tour across America. Trish kindly showed me the incredible palaeontology collections at the North Carolina Museum of Natural Sciences. Although having never met before in person, that shared connection through our museum work gave us an instant familiarity and we got on effortlessly. Trish kindly showed me the Pleistocene mammal collections, listening patiently as I chatted excitedly looking at enormous Ground Sloth bones. I asked about her favourite collections in the museum, and she quickly replied “our Ediacaran fossils”. I slowly put the Ground Sloth vertebra carefully down, and asked in case I misheard, “You have Ediacaran fossils in North Carolina?” Trish’s eyes sparkled. She walked me across the enormous store room, opened a cabinet, and without even pausing, pulled open the wooden drawer. There it was. It was beautiful.

For those readers who might read this as some unspoken love story, you would be right. This was a ménage à trois between a slab of 545 million year old rock and two people who had utter, complete, unconditional respect for it it. As Trish handed me the large piece of rock, I will admit, I didn’t see it straight away. That small dark smudge was it. And then I spotted more. These were fossils. Not of anything familiar, but of some colonial algae that lived long before the first trilobites: these were fossils from the very dawn of life. And both these fossils and Trish feature in this book.

The fossils I ware looking at were long thought to be burrows of some unknown little critter. Trish, along with Martin and a few more researchers, looked at these fossils with fresh eyes and discovered that these burrows were actually the remains of some strange colonial algae. Once growing on the 545 million year old sea floor, they were broken off and spread along the sediment by currents. That these burrows were not really burrows was an important finding because burrows in the Precambrian were rare: now they were even rarer. Why were burrows so rare 545 million years ago? I won’t spoil it for you, but instead we will jump forward in time, to a place you will be more familiar with: the Pleistocene.

It was great to see a lot of space given to the incredible burrows of the giant sloths and giant armadillos in South America. These creatures dug burrows. And they were huge.

The impressive underground burrow system of the giant ground sloth (Image reproduced with kind permission from Antony Martin)

We wrote about the enormous burrows of giant ground sloths, but Martin uses his knowledge to dig a little more than I could have. There are hundreds of massive tunnels in South America. Some have been filled in with sediment, while others lay undamaged. For some time they were thought to have been dug out by indigenous peoples, but there was very little evidence of human occupation in the tunnels. The smoothness to the tops of the tunnels suggested that something big was moving through these tunnels. And enormous claw marks in the sides of many of the tunnels indicates that this something very big was digging them. The claw marks give evidence to the makers: South American Pleistocene mega-fauna was impressive, but the claws of Giant Ground Sloths and Giant Armadillos matched the marks on the walls. No one knows why were they digging such huge burrows. Some had chambers at the end, indicating a nest. Burrows are used by many animals to bring up their young and protect them and baby Giant Ground Sloths would have been much more vulnerable than the adults.

The hand of yours truly on one claw from the North Carolina Giant Ground Sloth. This was just one claw. It was enormous!! (Photo by Jan Freedman. Taken at North Carolina Museum of Natural Sciences)

Sloths, Ediacaran fossils, dinosaurs, worms and the Gopher tortoise. These and many other examples of creatures both past and present open your eyes to how amazing life actually is. You know Martin loves burrows. It leaps from the page like the dirt kicked back by a gopher tortoise. Many people would find burrows, especially obscure fossil burrows, dull. Martin brings them to life, so much that you can visualise the creature making them as you read.

It was not because I am a total geology geek that I really enjoyed this book. It has everything you want in a popular science book: full of facts and information, well written, jokes to keep you on your toes, and stories to bring the pages alive. I loved Martin’s figures in the book and his non-standard scales for the images, which readers would easily relate to.

It was wonderful to see some fun figures in this book, including this Cave Bear den with a scale inspired by our very own Rey. (Image used with kind permission from Anthony Martin)

This was quite an achievement of writing: Martin covers the natural history of burrows over the last 550 million years giving some very well deserved air time for these normally ‘unsexy’ trace fossils. Not only will you discover about some quite fantastical creatures that once walked across the land, you will look at the world differently. So many animals rely on burrows for protection, warmth, mating, bringing up the young, or even simply using another animals burrow as their home. Many animals have actually survived disasters because they live in burrows.

Burrows are not seen, and so are often forgotten about. Next time you are on a walk have a look. Not just for animals scurrying about, but for the evidence they leave behind: foot prints, scat, or even a burrow.

Written by Jan Freedman (@JanFreedman)

Follow Anthony Martin on Twitter (@Ichnologist)

Follow Trish Weaver on Twitter (@somecephalopod)

Further Reading:

Martin, A. J. (2017). The Evolution Underground. Pegasus Books. [Book]

Posted in Giant Ground Sloth | Tagged , , , , , , , , , , , , | 2 Comments

No Bullwinkle

A fine specimen of a bull moose (Alces alces). Public Domain Image

A fine specimen of a bull moose (Alces alces). Public Domain Image

On Twilight Beasts you will hopefully have seen some of the amazing mummified animals of the Pleistocene. Some of these mummies may even be like old familiar friends to you: the woolly mammoths Lyuba, Buttercup, Khroma, Dima, Yuka, and Mascha, Sasha the woolly rhino, blue babe the steppe bison. Let’s not forget the recent discovery of my personal favourites Uyan and Dina the gorgeous cave lion cubs. Given the paucity of remains of extinct ice age mammals that have come to us from through the ages, the permafrost mummies that retain exquisite detail of hair, tissue, and skin allow us all to look into the face of a vanished time. You would think that any species we had almost complete mummified bodies from would be ice age superstars, with the level of pop-culture saturation that mammoths have attained. You’d think, but that’s certainly not the case. For instance, I’m willing to bet that few of you have heard of the stagmooses (stagmeeses? stagmice?), Cervalces scotti and Cervalces latifrons! Well, these guys are super-weird, and are represented in the fossil record by not one, not two, but three remarkably complete permafrost mummies.


Two Cervalces mummies from Alaska. Image taken from R.D.Guthrie’s classic “Frozen Fauna of the Mammoth Steppe”. I would dearly love to see a photo of these specimens but I have been unable to source any.

Some background: imagine if South Park style, a moose made sweet, sweet love to an Irish Elk and had some adorable babies. That’s essentially what a stag-moose looks like.

Cervalces scotti. Image by Stefano De Mateo/MUSE via Wikimedia Commons

Cervalces scotti. Image by Stefano De Mateo/MUSE via Wikimedia Commons

Another view of Cevalces scotti, at the Royal Ontario Museum. Image by Staka via Wikimedia COmmons

Another view of Cervalces scotti, at the Royal Ontario Museum. Image by Staka via Wikimedia Commons

C. scotti is the species known from North America, and C. latifrons the species known from Eurasia, but whether the two are actually close relatives or not nobody really knows (some refer to C. latifrons as Alces latifrons, moving it to the genus of the true moose, Alces alces). C. scotti may even be derived from Middle Pleistocene immigrants of C. latifrons. Whatever the relationship C. scotti definitely seems to be a genuinely distinct genus and species. Like Megaloceros, this species has a deer-like face and impressive rack, but in common with the moose, it has long legs and more palm than tines on its antlers. The antlers are themselves phenomenally complex features. Perhaps some of the most complicated headgear ever to evolve in cervidae and also fantastically variable throughout the species range.

Alces latifrons antlers. Aalen museum, image by Ghedoghedo via Wikimedia Commons

Alces latifrons antlers. Aalen museum, image by Ghedoghedo via Wikimedia Commons

Alces latifrons antlers,

Alces latifrons antlers, from the Natural History Museum of Mainz, image by Ghedoghedo via Wikimedia Commons


A.-C. Examples of Cervalces scotti E. Example of Cervalces latifrons from Churcher et al.

Cervalces scotti is something of an enigma. Despite the mummies (which all frustratingly lack the preserved head that would give us vital information on the character of this species) and some nearly-complete skeletons, pretty much everything else is up for discussion. Confined to northeastern America and Yukon/Alaska it appears to have had fairly specific habitat requirements, although what these were is ambiguous. Perhaps some kind of nonanalog environment was the preferred home for the stag-moose, something boggy and open, with not much woodland, which would explain why so many of their fossils are found in excavated peat layers. Whatever its requirements, radiocarbon dating shows that Cervalces scotti disappeared around 10,800 years ago. There is currently no evidence for human predation of Cervalces in the Americas, but bones of this taxon were found in the same layer as bone points at the archaeological site of Sheriden Cave in Ohio. It also is likely to have been affected by competition with the true moose (Alces alces) which came over the Bering land bridge about the same time.

It seems the stag-moose is an open book waiting for some dedicated researcher to step in and show an interest. The species complex is ripe for ancient DNA analyses that could resolve questions of paraphyly or stable isotope analyses to narrow down the vegetational needs of this unique lost species.

Written by Ross Barnett (@DeepFriedDNA)

Further Reading:

Churcher, C. S., and J. D. Pinsof. “Variation in the Antlers of North American Cervalces (Mammlia; Cervidae): Review of New and Previously Recorded Specimens.” Journal of Vertebrate Paleontology 7, no. 4 (1988): 373-97.[Abstract]

Guthrie, R. Dale. Frozen Fauna of the Mammoth Steppe: The Story of Blue Babe. London: The University of Chicago Press, 1990.[Book]

Long, C. A., and C. J. Yahnke. “End of the Pleistocene: Elk-Moose (Cervalces) and Caribou (Rangifer) in Wisconsin.” Journal of Mammalogy 92, no. 5 (2011): 1127-35.[FullText]

Redmond, B. G., and K. B. Tankersley. “Evidence of Early Paleoindian Bone Modification and Use at the Sheriden Cave Site.” American Antiquity 70, no. 3 (2005): 503-26.[Abstract]

Schubert, B. W., R. W. Graham, H. G. McDonald, E. C. Grimm, and T. W. Stafford. “Latest Pleistocene Palaeoecology of Jeffersons Ground Sloth (Megalonyx Jeffersonii) and Elk-Moose (Cervalces Scotti) in Northern Illinois.” Quaternary Research (2004).[Abstract]

Widga, C., T. L. Fulton, L. D. Martin, and B. Shapiro. “Homotherium Serum and Cervalces from the Great Lakes Region, USA: Geochronology. Morphology and Ancient DNA.” Boreas 41, no. 4 (2012): 546-56.[Full Text]

Posted in Extinction, Irish Elk, moose, Uncategorized | Tagged , , , , , , , , , , , , , , | 2 Comments