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

Battle of the Birds

I grew up in an area on the shoulder of a forested mountain, beside a series of lakes which are speckled with modern crannogs, replacements of ancient originals which are long gone. The shores and islets are rich in birdlife, as much as frog, fish and furry critters. A couple of years ago, the local paper ran a story of a hoodlum alpha male swan who was attacking small dogs, attempting a bit of Zeus and Leda-type action on other wildlife and unwary children, and apparently assassinating his love rivals. They shipped him off to someplace else, but his flailing wings and killer instinct caused quite a stir at the time. So much for the delicacy and grace of Swan Lake, eh?  Birds are tough cookies.

Waterfowl are a particular love of mine, which is probably why I find myself writing about odd birds (okay, that, and I’m an odd bird meself). I’ve paparrazi’d birds around the world on my peregrinations. I fell rather in love with the snowy white ibis picking through the riverside fields of the southern Nile: those long beaks extracting sustenance from the waters. More humans than me have been charmed by their otherworldliness – the ibis made its enchanting way into mythology as the avatar of the Egyptian god of wisdom Thoth, who chose avian form when he popped onto earth to observe human behaviour. Thoth was a proper sort of deity – no smiting or entrapment of poor mortals. Instead, Thoth was bookish, smart, all about the readin’, writin’ and ‘rithmatic; the Egyptian pantheon’s non-violent nerd.


The ibis-headed Egyptian god Thoth: much preferred books than boxing matches ( image from Pinterest,

Now, maybe they made ibises different in the Pleistocene, but there wasn’t much of bookish, scholarly Thoth about Xenicibis xympithecus, our utterly peculiar and amazing Twilight Beast. Roughly the size of a chicken, this extinct Jamaican ibis was unusual on two scores – it was one of only two known flightless ibis (the other one was in Hawaii…islands always have odd beasties), but it is the only bird known to have packed a punch, quite literally. For Xenicibis xympithecus had a unique wing structure which meant its wings were able to be used like a fist or a club.


From Encyclopaedia Britannica, the chicken sized slugger itself, Xenicibis xympithecus

The first fossils of this strange fisted ibis were found in the 1919- 1920 excavations of the Long Mile Cave complex in Jamaica, but it was only in 1976, when the bones were re-examined, that suspicions were raised about just how very unusual this flightless bird actually was. Other examples of the critter started popping up in Jamaican and American antiquarian collections, such as those found at Swansea Cave, also in Jamaica. Moral of this story, and many others is that those mouldy old collections from Colonial times beg for fresh eyes and new techniques; after all, just look at the bear patella discovered loitering in the National Museum of Ireland last year!

There had been a bit of a theory put forward in 1982 that the punchy little bird had also been present in Cuba , but the flightlessness aspect put paid to that. Xenicibis xympithecu was a Jamaican creature through and through, evolved to fit in with island life. Islands do funny things to species; Darwin thought of insular evolutions as being living, breathing products of evolutionary ‘laboratories’, with creatures adjusting to fit perfectly into their environmental niches. He didn’t perhaps realise just how swiftly that process can move – recent research has shown that island species can undergo accelerated evolutionary changes sometimes within decades, but certainly within several thousand years , a blink of an eye in the sequences of life on earth.

When more fossil evidence was added to the strange case of X. xympithecus  during the 1970s and 80s, a better understanding of the bird was created. All birds, flightless or not, have ‘arm’ bones just like our arms, comprising of radius and ulna, except of course much lighter to allow usually for flight. Those bones lead to the carpometacarpus, basically the equivalent of the human fist. The structure around and behind these bones was formed to allow the bird to literally swing a hefty punch with its wing. Instead of being rudimentary little pretend wings, with lightweight bones, this ibis was sturdy, strong and heavier boned.


Image from Longrich and Olsen 2011 showing the unique lower ‘wing’  ( labelled (h) ) of the Jamaican ibis

Some of their fossil bones actually have visible breaks in them, so it would appear these club-like wings were being used to thump something… but what? Could it be like the over-amorous killer swan of the Belfast hills who had to be deported to a sanctuary for delinquent ducks? Could the baseball-bat arms of X. xympithecus be used to slug out the best mate – survival of the species turned into Fight Club? It’s not unlikely, or at least were used as defence of some sort, but it may be more about territory than mating, as there really does not appear to be any sexual dimorphism in the bird bones. The Solitaire bird (also flightless, and also very extinct) of Rodrigues Island in the Indian Ocean had developed a bony knob structure on its wings, which it used to hammer its love rivals and win not just the lady pigeons but the best nesting areas. Extant ibises have been observed holding each other’s beaks and flapping their wings at each other furiously when enraged and either protecting territory or their young, so the amazing Xenicibis was very likely to use those mitts as well.

Depending on which version you believe, the birds died out between 10,000 and 2200 years ago. Its demise may have been caused by an environmental tilt in favour of one of the island’s predators, such as the yellow boa snake, or the extinct little monkey species Xenothrix mcgregori, as its bones were also found in situ at Long Mile Cave at the same excavation as our battling ibis. Nature is fragile –all it takes is a wobble in climate, a lurch of environmental change, and non-adaptive species vanish forever. Jamaica, like the entire Caribbean, has experienced climate shifts linked to changing tides and air currents as well as sea level changes throughout the Pleistocene and Quaternary periods. It also may not be surprising to know that Jamaica was only settled 2500 years ago by the Arawak peoples, who were migrating across the Antilles at that time.  Draw your own conclusions if you choose the more recent terminal date for Xenicibis.


Long Mile Cave, Jamaica. image from

So, there you have them – the amazing, strange and sadly extinct ibises with hurley-sticks for arms, who probably fought for territory and lurve with the same gusto. Some ibises round the Nile perhaps remember Thoth and his scholarly demeanour, as they glide on cloud-white wings among the hibiscus and palms of southern Egypt. But some, perhaps, remember when they were warriors. Birds, after all, have little dollops of dinosaur deep within their genes, and I like to think they remember that past, in a similar way to Mr Prosser in Hitch-Hikers’ Guide to the Galaxy , who remembered his bloodline to Genghis Khan at inopportune moments! We will never know if humans ever did witness the battles of the ibises amidst tropical rain forests, or seaside coves. But I’ll bet one thing – if they did, it must’ve been  quite a sight to behold the battle of the birds!.

Written by Rena Maguire (@JustRena)

Further reading:

Donovan, S.K. and Paul, C.R., 2011. ‘A diverse terrestrial fauna in the Pleistocene of Jamaica: the treasures of the Red Hills Road Cave’. Geology Today. 27.5  pp.173-180. [Full Article]

Donovan, S.K., Baalbergen, E., Ouwendijk, M., Paul, C.R. and van denHoek Ostende, L.W., 2013. ‘Review and prospectus of the Late Pleistocene fauna of the Red Hills Road Cave, Jamaica’. Cave and Karst Science. 40.  pp.79-86. [Full article]

Longrich, N.R. and Olson, S.L., 2011. ‘The bizarre wing of the Jamaican flightless ibis Xenicibis xympithecus: a unique vertebrate adaptation’. Proceedings of the Royal Society of London B: Biological Sciences. [Full article]

McFarlane, D. A., Lundberg, J., and Fincham, A. G. 2002. ‘A late Quaternary paleoecological record from caves of southern Jamaica, West Indies’. Journal of Cave and Karst Studies 64.2. pp 117-125. [Full article]

Millien, V., 2006. ‘Morphological evolution is accelerated among island mammals’. PLoS biology. 4.10. [Full article]

Morgan, G.S., 1993. ‘Quaternary land vertebrates of Jamaica’. Geological Society of America Memoirs. 182.  pp.417-442. [Abstract only]

Olson, S. L. and D. W. Steadman. 1977. ‘A new genus of flightless ibis (Threskiornithidae) and other fossil birds from cave deposits in Jamaica’. Proceedings of the Biological Society of Washington  90. pp 447–457.

Olson, S. L. and D. W. Steadman. 1979. ‘The humerus of Xenicibis, the extinct flightless ibis of Jamaica’. Proceedings of the Biological Society of Washington. 92 pp 23–27.

Olson, S. L. and A. Wetmore. 1976. Preliminary diagnoses of two extraordinary new genera of birds from Pleistocene deposits in the Hawaiian Islands. Proceedings of the Biological Society of Washington. 89. pp 247–258.

Scudder, S., 1991. ‘Early Arawak subsistence strategies on the south coast of Jamaica’. Proceedings of the Thirteenth International Congress for Caribbean Archaeology, Reports of the Archaeological-Anthropological Institute of the Netherlands Antilles .9.  pp. 297-315

Steel, L and Hume,J . 2013. ‘Fight club: a unique weapon in the wing of the solitaire, Pezophaps solitaria (Aves: Columbidae), an extinct flightless bird from Rodrigues, Mascarene Islands’. Biological Journal of the Linnean Society 110.1. pp 32–44. [Full article]

Suarez, W.  2001. ‘Deletion of the flightless ibis Xenicibis from the fossil record of Cuba. Caribbean Journal of Science. 37.1/2  pp.109-109. [Full article]

Posted in Ibis | Tagged , , , , , , , , , , | 3 Comments

Horned Armadillos and Rafting Monkeys

Walking back along the sandy path I could feel the refreshing cool air on my bare shoulders. The sun was starting to make its way down the horizon, turning the sky to fire. Rustling in the branches in the overhanging trees makes me stop. A little tuff-eared marmoset monkey sits confidently watching me. Then I spot another. More rustling and another appears. This well trodden sand path is the only way to one of the most beautiful beaches on Ilha Grande. These monkeys know it. Apple cores, biscuit remains and other food debris are evidence of countless tourists feeding these cute looking miniature monkeys. Suddenly, they scarper: as fast as they came, they vanish. A deep low growl rumbles in the trees behind. I knew there were jaguars in Brazil, but I hadn’t heard that jaguars had made it to the island. I wasn’t going to wait around to find out.

A gorgeous little Brazillian Common marmoset monkey. (

A gorgeous little Brazilian Common marmoset monkey. (Photo by Carmen Busko. Public Domain)

Brazil, and South America in general, has truly incredible wildlife. From Darwin’s Rhea to the wonderfully fun tapirs, this is a haven of wonderful creatures. It has an incredibly rich and unique fauna which has lived there over the last 65 million years. Why was it so special? What the heck is litoptern or a sabre toothed metatherian? To answer these questions, and more about the ancient mammals of South America, I was lucky enough to be offered a chance to review Horned Armadillos and Rafting Monkeys.

Such a concise, detailed book could only have been written by Darin Croft, at Case Western Reserve University. An expert in fossil mammals from South America, this is Croft’s magnum opus on his life’s passion. He wanted to put this book together to show off the incredible and weird animals that used to live there, most of which, he notes, people have never heard of.

South America is unique. Until about 3 million years ago, it was alone, drifting. After the break up of the mega-continent, Gondwana, the South American landmass split off from Antarctica. And here, for around 30 million years, it floated. Animals were pretty much isolated from the rest of the world, and evolved into their own unique forms. Croft takes the reader to meet these distinctive beasts from the very beginning of the Paleocene all the way up to the Quaternary.

The two major landmasses Laurasia in the north, and Gondwana in the South, around 200 years ago. (Image from here.)

The two major landmasses Laurasia in the north, and Gondwana in the South, around 200 million years ago. (Image Public Domain.)

Honestly, I wasn’t too sure when I first flicked through and saw the layout. First impression was that it looked like another textbook. As we all know, first impressions can be deceiving. As I began to read it, the layout made sense. It worked for the story Croft was crafting: the mammals that lived in South America over time.

Each chapter focuses on one palaeontology site looking at some of the key fossils found there. Each site is a unique piece used to create a detailed picture of the diverse animals in the past. From Bolivia to Argentina, we travel across the continent where digging the dirt allows us to read the pages in geological time. Something struck me as I read: by talking about the context the fossils were found in we go right back to the very roots of how we know what we know, and how there is still so much to learn. I liked this a lot! Looking at fossil sites (with some wonderful photos) reminds us that discoveries, either by serendipity or well planned, funded palaeontology digs, are the result of a lot of hard work. This is where the unglamorous real work begins out in remote places with determined, dedicated teams of people meticulously looking through the sediment for fossils. Often they return empty handed, or at best, with a few fragmentary fossils; tantalising hints of what could be.

Each site is well introduced with their place in the story of South America. It is clearly written for a general audience and although jam packed full of facts, Croft is gentle on the jargon. If you are wanting a personal touch, a little something to bring the author alive, you won’t find it past the prologue. But that actually doesn’t matter too much. You will find yourself lost in the incredible creatures filling the pages of this book. From giant carnivorous armadillos to tiny weird horse-like litopterns, this is a feast for the palaeo-naturalist.

A horned armadillo (Peltephilus ferox) carefully scouts around the entrance to its burrow before embarking on a search for food. Illustration by Velizar Simeonovski and provided courtesy of Darin Croft and Indiana University Press.

A horned armadillo (Peltephilus ferox) carefully scouts around the entrance to its burrow before embarking on a search for food. (Illustration by Velizar Simeonovski and provided courtesy of Darin Croft and Indiana University Press.)

Key species found at each site are listed, with some nice information about what it was and what the environment was like. Lovely artwork by Velizar Simeonovski means there is a visual connection with the extinct beasts and not just an unreadable Latin name on the page. For me the real gem of this book is the photographs of the fossils of each creature written about. Like the fossil sites mentioned above, this brings it all back to how we know what we know, and the hard work of the palaeontologists.

There are dozens of incredible creatures highlighted in this book. Our giant swimming sloth makes an appearance, as do several other Twilight Beasts. With nearly 30 million years of isolation, South America really was a unique ecosystem. Armadillos and sloths evolved there, and it is later in the Pliocene and Pleistocene where we see their giant forms. There was a very familiar looking sabre toothed relative of the marsupials, Thylacosmilus atrox which superficially looked like a cat, but was not even slightly related (the sabre toothed metatherian I mentioned earlier).

Later towards the end of the Pliocene, around 3 million years ago, North America and South America are joined by the Isthmus of Panama. South America was alone no more. This connection had an enormous impact on the ecosystems: lots of animals from both continents were able to migrate to new ecosystems in what is known as the Great American Biotic Interchange (GABI).

Those South American animals that migrate north were not as successful as the northern species that moved south. The giant sloths and glyptodonts only made it into lower north America because the environment was much drier than they were used to. The newcomers to the south, however, thrived in the tropical environments. Tapirs, animals almost symbolic of the Amazon, originated in North America. Rabbits, pikas, bears, weasels, and deer (including the smallest deer species, the gorgeous Pudu) are all animals that made their way south, and adapted so well they are well and truly at home. The only northern Order which didnt survive until today were the Proboscidea, the Gompotheres that made it south became extinct around 10,000 years ago.

The Great American Interchange showing the creatures that moved to explore new lands.

The Great American Biotic Interchange showing the creatures that moved to explore new lands. The green animals were South American beasts that travelled north. The blue animals are North American beasts that moved south. (Image Public Domain)

It may be my geeky side coming out, but I enjoyed it. It has a lot of information, which can be pretty heavy going at times, but I know I will be going back to this book again and again. It also answered my ponderings about cats and monkeys. Cats too made it into South America after the Great American Biotic Interchange. Monkeys arrived in South America way before the cats arrived: between 37-40 million years ago, some African monkeys came adrift in the Atlantic ocean and landed in South America (this probably happened several times, with many rafting monkeys not surviving).

Hop into a time machine, and land around South America between 65 million years ago and today, and this book would be your guide.

Written by: Jan Freedman (@JanFreedman)

Further Reading:

Croft, D. A. (2017). ‘Horned armadillos and Rafting Monkeys: The fascinating fossil mammals of South America.’ Indiana University Press. [Book]

Read more about the wonderful Gomphotheres here.

Love the Giant Armadillo, find out more here.

Charles Darwin’s lost giant ground sloth – here.

More about the enormous giant ground sloth, Megatherium – here.

Did you know giant ground sloths dug burrows? Here you are.

The worlds smallest deer – here.

Posted in Giant Ground Sloth, Gompothere, Ground Sloth, Macrauchenia, Pudu, Thalassocnus | 2 Comments

Open your eyes and see beauty

“Open your eyes and see what you can with them before they close forever.”  Anthony Doerr, 2015.

She is beautiful: She is quite possibly the most beautiful thing in the room. I have travelled to America to see her, and she does not disappoint for the one day I get. I stand there looking at her from afar. She is subtle. She is slight. She does nothing to stand out. She is there but almost isn’t. She just is.

I watch as several people walk right past her: their eyes on something else. A quick glance to their left and they would see her. I will them to look left; with all my grey matter, I will them. I almost shout out ‘look at her!’ but then one person does turn: they look but they don’t see.

Often the most beautiful things are right in front of us, and we don’t see them. Maybe we don’t see because we are not looking for it. Or maybe we don’t see the beauty in front of us because our minds are often removed from it: the need to look for bigger things, the safety of seeing familiar things. Our heads are so often full of other, more mundane thoughts that we cannot focus. Ironically, to clear your head and notice the detail of what is really in front of you relaxes you more: other thoughts are gone, worries no longer there, your brain is taking in the colours, the patterns, the detail. And it feels wonderful.

I stood in that room, watching people walk by, and was quietly sad because less than a handful stopped to look at her. I would go and see her every day if that room wasn’t around 4000 miles away from where I sit now.

And her. Just standing there. Just looking so beautiful. I still remember all the detail: those big eyes, the curve of her body, the smoothness of her skin. She was the most beautiful creature I had ever seen.

I was so lucky to see her. Even so, she wasn’t really there. I missed her. By a mere 11,000 years.

The enormous Columbian Mammoth on display at La Brea, pulls the visitors away from other displays. Author for scale. (Image, authors own.)

The enormous Columbian Mammoth on display at La Brea, pulls the visitors away from other displays. Author for scale. (Image, authors own.)

La Brea Tar Pits Museum, in Los Angeles, is the last place you’d expect to see her. But then again, this is an incredibly rich site; home to over 650 different species of Late Pleistocene fauna. You are greeted by a cleverly lit, fully mounted skeleton of a Giant Ground Sloth, (Glossotherium harlani) that looks alive although it is just bones. The lights are low, giving a quiet, subdued ambience. As you walk into the first main room, to the left of there are displays flat against the wall, which at a glance look like some bones of some small creatures, lots of 1970s font, and a model of some deer. To the right, in the centre of the room, is where the big specimens are: the specimens people want to see, the sexy specimens, the specimens people gasp at: the Mastodon, the enormous camel (Camelops), the goliath Columbian Mammoth. Following this trail of breadcrumbs leads you straight out into the next room, of giant birds and Dire Wolves. Break away from the trail, however, and there are wonderful displays against the wall: teeth from the Imperial Mammoth, bones from the ancient Bison, and her, the beautiful dwarf pronghorn, Capromeryx minor.

A display you could just walk past. But take a moment to stop. And appreciate how amazing this creature was. (Photo by Jan Freedman)

A display you could just walk past. But take a moment to stop. And appreciate how amazing this creature was. (Photo by Jan Freedman)

I see how easy it is to walk past her. She just looks like a small deer. Nothing special. Nothing compared to the enormity of the Columbian Mammoth, or the terror of the sabre-tooth cat. That is of course, if you don’t look. Open your eyes and you will see not a little deer, but something much rarer. Something much more beautiful.

The dwarf pronghorn belongs to an almost vanished group, the Antilocapridae, where the only surviving species is the Pronghorn Antelope in North America. Superficially looking like antelopes, they were not. They are actually more closely related to the ancient group of Giraffes (another beautiful group which once held a huge array of magnificent species, but is now down to just a couple today). Just as in Giraffes, the horns on pronghorns are bone growths out of their head – unlike deer where the antlers are not true bone.

There were once over a dozen different pronghorn species leaping across the North American wilderness, with their long, elegant legs. One species would have been at home more in the forests than the open plains: our Dwarf Pronghorn. And she was a small species, the smallest species of pronghorn: perhaps reaching up to my hips at just 60cm tall at the shoulders.

The Dwarf Pronghorn was the smallest pronghorn species. (Image by Jan Freedman)

The Dwarf Pronghorn was the smallest pronghorn species. (Image by Jan Freedman)

Capromeryx minor was a relative late comer to the Pleistocene scene. Although the Genus Capromeryx evolved around 5 million years ago, our little friend arrives just 300,000 years ago. Fossil finds show the distribution was fairly restricted to southern North America, perhaps an indication of their preferred habitat: more wooded than open, like the tiny pudú. In almost 300,000 years, they got smaller and smaller.

When you look at the skeleton of this animal, you can see how stunning it is. Long, slender legs hint at quick bursts of speed in an attempt to escape from predators. (I like to imagine them leaping away instead of running.) They had very tough teeth, indicating eating tougher plants than just grasses alone. Like all pronghorns Capromeryx minor had horns on their head, four in total: two slightly larger ones at the back, and two smaller ones at the front. Because they are extinct, we know so little about their lives: It may be that the horns used by males for fighting each other, because the males were larger.

These gorgeous little animals almost made it through to today, with the last known fossils dating to around 11,000 years ago, at the end of the last major glaciation. We don’t know why they vanished. At the same time many other of the familiar beasts of the very Late Pleistocene were disappearing. Climate was warming, glaciers were melting, and humans were hunting. It is unlikely that humans hunted the dwarf pronghorn to extinction. But there could have been an indirect impact. Ecosystems are complex, subtle, but reliant interactions of food, temperature, water, animals and plants. At the end of the Pleistocene many of the large fauna were removed from the ecosystem, plants retreated and new ones took root, and temperatures changed fast. Our little beast couldn’t leap fast enough to keep up.

Life is incredible. It is so fragile, and yet can appear so robust. All of these animals that are not here today show us what the world was once like. It was, and still is, astonishing. It is not just the big, or the familiar which make our planet beautiful. Stop and look at it all: the fly on the windowsill, the bird in your back garden, the person sitting next to you. Life is beautiful. When you open your eyes, you can really see that beauty.

Written by Jan Freedman (@JanFreedman)

Further Reading:

David, E. (2007). Family Antilocapridae. In D. Prothero, S. Foss (eds). ‘The Evolution of Artiodactyls’. The Johns Hopkins University Press, Baltimore, Maryland. [Book]

Doerr, A. 2015. ‘All the light we cannot see.’ Fourth Estate, London. [Book]

Hernández Fernández, M. & Vbra, E. (2005). ‘A complete estimate of the phylogenetic relationships in Ruminantia: a date species-level supertree of extant ruminants’. Biological Reviews. 80:pp.269-302. [Full article]

Jimenéz-Hidalgo, E., O. Carranza-Castañeda, M. & Montellano-Ballesteros (2004). ‘A Pliocene record of Capromeryx (Mammalia: Antilocapridae) in Mexico’. Journal of Paleontology. 78(6): pp.1179-1186. [Abstract only]

Kurtén, B., & E. Anderson (1980). Pleistocene mammals of North America. Columbia University Press. New York. [Book]

Janis, C. & E. Manning. (1998). Antilocapridae. In: C. Janis, K. Scott, L Jacobs, (eds.), Evolution of Tertiary Mammals of North America, Vol. 1. Terrestrial carnivores, ungulates and ungulate-like mammals, Cambridge University Press, Cambridge, UK. [Book]

Martin, P. S. (2007), Twilight of the Mammoths. University of California Press. [Book]

McMenamin, M.A.S.; et al. (1982). “Amino acid geochemistry of fossil bones from the Rancho La Brea Asphalt Deposit, California”. Quaternary Research18 (2): 174–83. [Abstract only]

Morgan, J., & N. Morgan. (1995). A new species of Capromeryx (Mammalia: Artiodactyla) from the Taunton Local Fauna of Washington, and the correlation with other Blancan faunas of Washington and Idaho. Journal of Vertebrate Palaeontology. 15: pp.160-170. [Abstract only]

Murray, L. (2006). The smaller Artiodactyls: peccaries, oxen, deer, and pronghorns. Jefferson, G. and L. Lindsay (eds) In: Fossil Treasures of the Anza-Borrego Desert: the Last Seven Million Years. Sunbelt Publications, San Diego, California. [Book]

Stock, C., 1930. ‘Rancho La Brea. A record of Pleistocene life in California,’ Science Series. No 37. Natural History Museum of Los Angeles.

Posted in Columbian Mammoth, Dwarf pronghorn | Tagged , , , , , , , , , , , , , , , | 3 Comments

The most (and least) read blog posts of 2016

A huge thank you to all our readers for another wonderful year of Twilight Beasts! For some reason we didn’t receive our annual ‘summary of’ breakdown, so we shall pull it together for you!

Thank you for all your continued support and feedback on our blog post about these wonderful creatures of the past. Living in this virtual world, it is not that easy to show how much we appreciate your support with the blog. We hope that readers do get that from our tweets and our writing.


So, the eagerly awaited ‘best of’ list is here. Here are the most read (and the least read) beast blog posts of 2016. Show them a little love, and delve in deeper to discover more!

Bottom 5 read blog posts:

  1. Down the rabbit hole. Small islands and giant dormice take centre stage in this post looking at the fragility of species.
  2. The ancients of the forest. Not all our blog posts are about giant mammals. Or lizards. Or insects. This one is about a giant tree from New Zealand!
  3. T’was the night before Christmas. I guess, in the words of Pierce Bronsan, Christmas only comes once a year. But you can find out all about how Reindeers once travelled in their thousands across Europe!
  4. Baby’s got quack. Well, Japan once had a pretty big Duck. And this blog post has a pretty great title too.
  5. A tiny Twilight Beast in a world of giants. The least read post of 2016.: this deserves a click if only for the cuteness. Discover how the smallest deer, the Pudu, survived the last Ice Age.
Intricately carved from fragile mammoth tusk, these two reindeer look as though they are swimming, although this may be due to the lack of depth of a mammoth tusk. (image from the British Museum)

This beautiful object was carved from fragile mammoth tusk. These two reindeer look as though they are swimming, although this may be due to the lack of depth of a mammoth tusk. (Image from the British Museum)


Our top 5 read blog posts of 2016:

  1. Going Underground. A spectacular insight in to the lives of Giant Ground Sloths: some species dug enormous burrows!!
  2. The bear from Clare – new evidence for an early human presence in Late Pleistocene Ireland. Early last year a bear knee cap was discovered in the National Museum Ireland, with cut marks on it. Discover how this incredible find pushes back the date for humans in Ireland.
  3. Lost as the Moa is lost! The enormous flightless bird from New Zealand, the Moa, only became extinct around 600 years ago.
  4. On the Origins of our species. Our first guest blog post makes it in the tope 5! Find out where we come from, and how really, despite geography, we are all more closely related than we think.
  5. The hyena that was overshadowed by a Tyrannosaurus rex. Yay! A hyena post made it in the top 5!! This is all about the only hyena species that made it to America, he who saw the canon.
One enormous claw from a Giant Ground Sloth on display at North Carolina Museum of Natural Sciences. This enormous claw could dig through rock. To make tunnels!

One enormous claw from a Giant Ground Sloth on display at North Carolina Museum of Natural Sciences. This enormous claw could dig through rock. To make tunnels! (Image by Jan Freedman)

We hope you enjoyed our blog posts from 2016! We shall be bringing lots more new and exciting beasts to you for 2017.

Until then, enjoy the top (and bottom) 5 posts.

We wish all our readers a very happy and healthy 2017. Ross (@DeepFriedDNA), Rena (@JustRena) and Jan (@JanFreedman)

Posted in Brown Bear, Celebrate!, Giant Ground Sloth, Giant Maltese Dormice, Homo sapiens, Hyena, Kauri tree, moa, Pudu, Reindeer | Tagged , , , , , , , , , | 1 Comment

The Lost Sheep

I don’t know about you, but I love the spirit of farmers during lambing season. Coming from a farming background, I know the farmers’ mingled emotions of dread and hope and joy as the wee lambs are born often in the dead of winter. The late nights, camaraderie, the dance of life and death, sometimes played out against mountain snow; this time connects us with the farmers of prehistory. It is, above all, a time of hope when lambs are like tiny fuzzy reminders of springtime ahead and the renewal of life. I’m pretty sure even our pre-Neolithic ancestors gazed in admiration at the tiny, resilient animals frisking amidst the herds which passed through European refugia.

For this Twilight Beast, Megalovis latifrons, the giant sheep, we shall go back a little earlier than usual for this blog. Right to the very beginning, on the boundary of the Pliocene and Pleistocene. It was all happening during the Epivillefranchian period, that junction between the Pliocene and the earliest Pleistocene, about a 2. 6 million years ago. It’s believed there were numerous and regular large tectonic events, setting off a chain reaction of natural changes to the planet. Around this time, some species of hominins had made their way to Eurasia, and were also settling across Europe, though would get driven back by the gradual onset of glaciation through the Pleistocene. In Europe and Asia, there were numerous ‘dispersal’ events, occurring over a relatively short period of time, when whole faunal assemblages were replaced, and new critters arrived from other continents – elephants, horses and the rather charmingly named ‘Wolf  event’ of around 1.7 million years ago. Climate shifts were noticeable from the stable isotope analysis we have gained from ice cores. It was getting cold. Snow would fall, deep, and crisp and even….a dangerous time for sheep, a time when they can get lost.

The giant sheep

The giant sheep Megalovis latifrons, with our Rey for scale. This was an extremely large sheep.

There were only desperate hunter gatherers inhabiting Europe and Eurasia at this time, clinging to existence daily, battling everything the climate or lack of game could throw at them, so no shepherds watched flocks by night, or day– the Neolithic was quite a way yet!  This lack of attendance was probably quite welcome, or at least did not bother the giant sheep, Megalovis latifrons. There are very few fossil assemblages of this creature, although it is believed to have been around the size of a modern musk ox, with large, straight horns facing outwards like a sheep, and with similar dentition as well. There are some very early faunal assemblages from Verchets in Romania, indicating that Megalovis pre-dated the shift into the cold of the Epivillefranchian, and a substantial bone assemblage in Trlica, Montenegro dating right up to the early Pleistocene around 950,000years ago.


Horns of subspecies Megalovis balcanicus from CRÉGUT-BONNOURE and DIMITRIJEVIĆ 2006, reference in blog bibliography

Horns of subspecies Megalovis balcanicus (from CRÉGUT-BONNOURE and DIMITRIJEVIĆ 2006, reference in Further Reading)

This was the sort of sheep which could have delivered a bit of a lamb chop if it was provoked. It was a powerful and steady grazer of grassy open lands, but nimble enough to navigate the mountain ridges of Europe and Eurasia. We are not even terribly sure what they looked like, but we know from especially the Trlica assemblages they had pointy premaxillar bones on their jaws, which have been considered indications that they were fussy feeders, grazing on one single type of food, probably leaf-based. As folivorous creatures, chewing down cellulose-heavy leaves, they may have had slower metabolisms than modern sheep or goats. There’s been such a lack of understanding of this rare breed that palaeo folks weren’t exactly sure how to categorise them for quite some time – Sheep? Goat? Antelope? There is still sporadic new work on variations of giddy goats and sheep which may, or may not, be related to Megalovis.  There have been other branches of Megalovis identified, such as Makapania sp, which lived in Africa, although they may have been much more antelope-like than their European cousins.

We now have a pretty good idea that Megalovis was one of 19 species of Ovibovini, much closer to modern musk oxen and gorals than sheep as we understand them. The giant sheep was, to all appearances, a successful Pliocene species, yet by the middle of the Pleistocene, they were all extinct, which probably accounts for why there are no images of them (to date anyway) incorporated into cave art displays. And they must have been quite a sight, herds of extremely large sheep wandering the chilly grasslands of Europe. Until a complete assemblage is discovered, we still need to use guesswork about how they looked – though they probably resembled the extant goral – only larger.

A modern day Goral. (Image by FunkMonk on Wiki)

A modern day Goral. (Image by FunkMonk on Wiki)

So, if humans didn’t get the chance to over hunt Megalovis, what took the great sheep from the gene pool? As we mentioned earlier, the creatures were picky eaters (and feel free to use this cautionary tale on kids not eating their Brussels sprouts this Christmas!!). As vegetation changed with the creeping cold of the Pleistocene, the woolly jumpers just didn’t have enough to eat. Sadly, one of the perils of being a niche feeder is potential extinction should anything endanger the food supply. Gradually, they faded from the landscape, replaced by the less fussy Soergelia elisabethae and Praeovibos priscus. It would be the middle of the Pleistocene before species which we would well recognise today such as the ibex, would become the inheritors of the gap left by Megalovis.

We can only imagine how adorable the little lambs of the species must have been, and it is regretful that there are no known images of the creatures, compared to depictions of ibex, bison, mammoths and lions, which grace the walls of caverns across Europe, with their observations on nature and hunting, daubed skilfully by torchlight with charcoals and faith to express beliefs long since forgotten. If anything, our blogs echo the love of these animals from the deep past, our paintings made with pixels and words, communicating the same wonder as ancient peoples at this beautiful, amazing planet we live on.

One recurring theme of our blogs is how all species –ourselves included – are woven together in the warp and weft of climate and environmental change. Sometimes natural changes occur without human intervention, causing extinctions like Megalovis, replacing these creatures with more versatile species which can survive adverse conditions better. But life does go on… the miracle of adaption, rebirth and survival continues, just like modern little lambs at the worst time of the year may as well say ‘come at me bruh’ to wind and rain and snow. This Christmas, Hannukah, Kwanzai, or Yule (or simply winter break) we at Twilight Beasts wish you tenacity,  hope, courage and joy in these changing and often scary times.

And please – don’t be picky eaters – we want to share more amazing creatures with you during 2017!

From the Beast team of Jan, Ross and Rena – we wish ewe a wonderful season!

Written by Rena Maguire (@JustRena)

Further reading:

Bechev, D. and Georgiev, D., Paleobiodiversity of the Vrachanska Planina Mountains in the Villafranchian: a case study of the Varshets (Dolno Ozirovo) Early Pleistocene locality of fossil fauna and flora. [Full article]

Brugal, J.P. and Croitor, R., 2007. Evolution, ecology and biochronology of herbivore associations in Europe during the last 3 million years. Quaternaire. Revue de l’Association française pour l’étude du Quaternaire, 18. 2.  pp.129-152.

Cregut-Bonnoure, E, 2004. European Caprinae ( Ovibovini, Caprini) from the Plio-Pleistocene Boundary: a new interpretation. 18th International Senckenberg Conference Proceedings. [Full article]

Cregut-Bonnoure, E. and Dimitrijevic, V., 2006. Megalovis balcanicus sp. nov. and Soergelia intermedia sp. nov.(Mammalia, Bovidae, Caprinae), new Ovibovini from the Early Pleistocene of Europe. Revue de Paléobiologie. 25.2. p.723.

Gentry, A.W., 2000. An Ovibovine (Mammalia, Bovidae) from the Neogene of Stratzing, Austria. Annalen des Naturhistorischen Museums in Wien. Serie A für Mineralogie und Petrographie, Geologie und Paläontologie, Anthropologie und Prähistorie, pp.189-199. [Full article]

Gentry, A.W., 1970. Revised classification for Makapania broomi Wells and Cooke (Bovidae, Mammalia). Palaeontologia Africana. 13. [Full article]

Kahlke, R.D., García, N., Kostopoulos, D.S., Lacombat, F., Lister, A.M., Mazza, P.P., Spassov, N. and Titov, V.V., 2011. Western Palaearctic palaeoenvironmental conditions during the Early and early Middle Pleistocene inferred from large mammal communities, and implications for hominin dispersal in Europe. Quaternary Science Reviews. 30.11 pp.1368-1395.

Lehmann, U. 1953. Eine Villafranchiano-Fauna von der Erpfinger Hohle (Schwabische Alb). Neues Jahrbuch für Geologie und Paläontologie Monatshefte 10. pp 437-464

Lordkipanidze, D., Jashashvili, T., Vekua, A., Ponce de León, M.S., Zollikofer, C.P.E., Rightmire, G.P., Pontzer, H., Ferring, R., Oms, O., Tappen, M., Bukhsianidze, M., Agusti, J., Kahlke, R., Kiladze, G., Martinez-Navarro, B., Mouskhelishvili, A., Nioradze, and  M., Rook, L., 2007. Postcranial evidence from early Homo from Dmanisi, Georgia. Nature 449, pp 305-310.

Masini, F., Palombo, M.R. and Rozzi, R., 2013. A reappraisal of the early to middle pleistocene Italian bovidae. Quaternary International, 288, pp.45-62.

McClymont, E.L., Sosdian, S.M., Rosell-Melé, A. and Rosenthal, Y., 2013. Pleistocene sea-surface temperature evolution: Early cooling, delayed glacial intensification, and implications for the mid-Pleistocene climate transition. Earth-Science Reviews, 123, pp.173-193. Available at:

Spassov, N. and Crégut-Bonnoure, É., 1999. First data on the Villafranchian Bovidae of Bulgaria. Comptes Rendus de l’Academie des Sciences Series IIA Earth and Planetary Science7. 328.   pp.493-498.

Turner, A., 1992. Large carnivores and earliest European hominids: changing determinants of resource availability during the Lower and Middle Pleistocene. Journal of Human Evolution22. 2.  pp.109-126.

Posted in Giant Sheep | Tagged , , , , , , | 1 Comment