Bones and A Brain

Approximately 500 individual bones have been assigned to Ampelosaurus atacis from the excavation sites in Southern France. The femora from that location number approximately 27 individual specimens. These bones are not assumed to have come from singular individuals specifically, meaning that any number from 14 to 27 individuals may be represented by these femora. More interesting, however, is a non-specifically assigned braincase described in 2013 by Knoll, et al. The braincase includes the majority of the skull surrounding the braincase (this includes paired frontals, parietals with extensions, basioccipital, occipital, basisphenoid, parabasisphenoid, prootic, and laterosphenoid). The CT scans and 3D renderings from the description and research are available free on PLOS. The images from the study are wonderfully detailed and freely available, however, they are too large to upload. In order to look at them and appreciate them the link is contained in the reference below.

Knoll F, Ridgely RC, Ortega F, Sanz JL, Witmer LM (2013) Neurocranial Osteology and Neuroanatomy of a Late Cretaceous Titanosaurian Sauropod from Spain (Ampelosaurus sp.). PLoS ONE 8(1): e54991. doi:10.1371/journal.pone.0054991

Originals and Osteologicals

Le Loeuff's original description of the titanosaurid Ampelosaurus is available online as a PDF from the translators over at the Polyglot Paleontologist. The translation is well done and the paper is short but informative. Lacking from this version though are the figure plates and details to which Le Loeuff refers in the paper. The images are available online in other places, including the Academia page of the article uploaded by Le Loeuff. The osteology of the dinosaur has also been discussed in the book Thunder-lizards: The Sauropodomorph Dinosaurs. Some of the discussion for the topic is missing, but if the entire book is available it is worth borrowing and reading the entire section. If not, making do with the missing pages is not the end of the world either.

Video Reviews

Today I thought that we could try something a little different. There is not a documentary or news story or any other such video as we prefer to find on Mondays, however, there are a pair of reviews of the same Ampelosaurus model that we can view. One of the reviewers promises a discussion of the life history of the animal while the other only promises a review of the model. Either way a history of the animal must be discussed to review the authenticity of the model. I shall leave it to the videos to do the rest of the explaining for themselves though.

Tiny Titanosaurs for Tots

The sites that discuss Ampelosaurus are ample (pun intended). About sticks out first because it comes about alphabetically first and because it does, as usual, a very good job of summarizing the key facts and has been written up into an easy to read paragraph also. The NMH of London also has a site dedicated to the dinosaur, but puts out a lesser fount of information than the About page. This is okay to a point because they do make it up a little with a nice quality original illustration. It is not the same of course, as having quality information. KidsDinos and Prehistoric Wildlife both have respectable fact files and nice illustrations of the dinosaur. The search can continue beyond those sites, but the quality of the sites falls off quite a bit. There is no quality coloring page available today either, but these sites should keep young readers busy for a while and the illustrations can easily be used as references to draw their own dinosaurs today!

Protection and Little Titanosaurs

Compilation uncredited, dinosaur illustration is the same as yesterday (Dmitry Bogdanov).

Ampelosaurus, despite being a titanosaurid, appears to have been a somewhat smaller sauropod than is often thought of when we hear the name "titanosaur." This small-ish appearance of Ampelosaurus could be an artifact of the size of the femora discovered, but considering the number and diverse amount of remains that have been discovered, this most likely simply was a smaller titanosaur. This idea is probably best exemplified in illustrations (other than in the size comparison graphic above) by Alain Beneteau's depiction of a Taracosaurus attacking an Ampelosaurus. The head of the sauropod is, in all the illustrations that I have seen, are nearly identical. There is a known braincase and partial skull that accounts for the similarity in this aspect of all the illustrations. The back of Ampelosaurus is covered in osteoderms that have also been unearthed. As far as I know there is only a single photographed osteoderm available on the internet for our viewing.

Femora of France

©Dmitry Bogdanov

Discovered in France and Spain, Ampelosaurus atacis Le Loeuffe 1995 (Vine lizard of the Aude River) was a Cretaceous titanosaurid that presumably lived throughout Europe (though this has not been substantiated as yet). Many partial sets of remains of this dinosaur have been recovered, but the majority of those remains are actually represented by a single bone of the skeleton; the femur. The catalog of animals found near this dinosaur actually outweighs its gargantuan remains, but we can discuss that at a later time this week. Though known as a French dinosaur originally, some of the most interesting remains, including a braincase, have been discovered in Spain.

Human Ankle, Tarsier Leg

The morphology of Archicebus indicates that it is one of, if not exactly the last common ancestor of all primates from humans to monkeys. The largest similarities between primates and Archicebus are found in the metatarsals and the ankle, particularly the calcaneus. These bones are similar in shape and proportion to those seen in later primates. Archicebus, however, also possesses a skull, teeth, and limb bones that are much more common to a group of lower mammals (tarsiers).

Paper Time

The find of Archicebus has been important and monumental. The papers, many of the illustrations of which we have seen here this week, that have been published are few in number but are pretty informative and useful. The original paper and a completely independent paper that can probably be read in full with institutional access (I do not have that at home). Either way, the original paper is worth reading and it is the kind of paper that I never want to spoil by summarizing here. The discussion of haplorhine evolution is short, considering the overall paper length, but is definitely helpful in knowing where these tiny little primates rest on the tree of life.

News and Movies

To start, there is good news in the world. You can find that news here. It is very nice to be a part of the group and it is going to be quite a busy and fun time in the lab now.

In terms of video, there is a surprising lack of news stories associated with the relatively newly announced animal. The video shown here is a small snippet of that news. A slightly better news story can still be found on the NTD (New Tang Dynasty) television service website (the site states they are based in New York and broadcasts directly to mainland China and worldwide). This news story at least shows a portion of an interview with Ni Xijun that I saw the other day. Unfortunately, that interview was not in English at all. The NTD story translates it for us though thankfully.

Short Listed

Today there is not much out there. The newest finds usually have the fewest links and Archicebus is not that much different from any other fossil animal. There is a short round up on the About (http://dinosaurs.about.com/od/mesozoicmammals/p/Archicebus.htm) pages, which have always been pretty extensive in their coverage of the animal kingdom. Unfortunately, today is one of those rare days where we do not have a lot of specialized articles for people to let their kids loose on the internet to read. In that vein, however, it may be worth the time to go about searching safely and discuss what makes websites acceptable sources of information. There's a cheat sheet for that in case it is a topic that has never been discussed in your house (http://bit.ly/1CiN8tr).

Creepy Smiles

© Xijun Ni

The creepy smile of Archicebus achilles in this restoration is a little odd, though if I grabbed an insect that size for dinner as a rather small primate I would be just as happy as Archicebus appears to be. The rather large toes of the hindlimb in the reconstruction are true to those of the fossil remains as well. The nearly opposable thumbs on the forelimb are obviously useful for grasping insects like this rather large piece of dinner. The only thing that has not been alluded to with the fossil used as a model for the illustration is the potential ability of the tail to be used in a prehensile manner. The illustration does appear rather tarsier-like, which is a good thing as the animal is hypothesized to have been a member of the tree living between tarsier and monkey families.

That Prehensile Tail

© Xijun Ni

The first primates are probably what the populace thinks of when one says something like "prominent early mammals". Therefore, as the last of the strictly mammal weeks here, I present to allthe earliest recognized primate, Archicebus achilles Ni et al 2013. This little tree-scrambling primate is considered the first "haplorhine" or dry-nosed primate, a group that monkeys, apes, and humans all belong to as well. Its discovery in Asia supports hypotheses that primates first began to evolve in Asia and later migrated toward and eventually into Africa, where it is known that humans later evolved and originally migrated from. The name of the animal originated from the Greek and Latin combination meaning "Beginning monkey" with a reference to Achilles, supposedly due to a significantly novel calcaneus discovered in the fossil. The animal is comparable in weight to the smallest extant primates (Mouse Lemurs) but is thought to have been active during the day rather than nocturnally, like most small mammals were, and still largely are, when this little creature first began to confidently stride around on the planet.

Cartoon Popularity

Cimolestes and other small mammals do not make it into the toy market very often, which is quite okay. They often do not make much of a mark in the video game market either. Cimolestes does, however, actually hit a good chunk of the popular arena marketed toward kids thanks to Dinosaur Train that includes both video games, even if it is still just a small portion of the games online that are devoted directly to the Dinosaur Train genre. The same goes for toys; Cimolestes only really appears where Dinosaur Train characters are concerned. These are both hard to find online either way though. It actually seems that the only way that Cimolestes has been introduced to the public over the years is through technical literature (popularized books are absent) and the Dinosaur Train series.

What About Your Tail?

I noticed that there was a pretty fantastic piece of art out there but it has no illustrator credit and no dollar amount attached to it. Therefore, I present it to you today as a link to the bureau that apparently manages its release. I do not think that breaks any kind of copyright rules, so it should be okay. The image does not really address what I wanted to address today, but it is pretty fantastic looking. The thing I really wanted to look at today is the tail of Cimolestes. Most small rodents (e.g. anything smaller than Capybara and Beaver) we think of either have wiry little tails and use them as balancing tools or big bushy tails that can be used as balancing tools or to help provide warmth. Cimolestes is usually depicted with the wiry looking tail that we see in mice, rats, and shrews. It stands to reason, and considering the environment at the time, that there was not much need for Cimolestes to have a bushy tail for warming itself if this wiry tail counterbalance is correct. Balancing as it scrambled up and down the tree was probably the limit of what Cimolestes needed its tail for anyway considering that it used all four limbs to scramble around the tree. Any small help in balancing while running was a benefit for an animal scrambling away from dinosaur that could swallow it whole if it tripped or otherwise lost its balance when it was running away. Long tails in extant mammals sometimes serve the exact same purpose. Think about how well adapted Cimolestes was the next time you see a rat, mouse, or even a cat running along a fence top!

Not Necessarily Correct

©Carl Buell

The image presented here is from a news story about a small mammal that mentions Cimolestes and discusses their habitat. The story does not mention whether or not this image of Cimolestes or the other small mammal that is being discussed, so we can use this image today to go with the other papers and articles that I found about Cimolestes that are pretty interesting. The first article that was really good today actually discussed Cimolestes in a round about way as it actually centered on the evolution of placental mammals rather than the mammal itself. Woodward's 1910 description of a mandible found in Montana is also a good addition to the reading list for today. It notes how small details can be used to attribute something as small as a mandible to a known species. The final paper worth reading for the day is about two new potential specimens from the Gobi Desert. The specimens do not belong to the genus Cimolestes, but that animal is described and discussed in comparisons to the two genera that are being described in the paper. This kind of cross identification and description are important to be aware of and to be able to understand when reading future papers (consider it a future preparation tool for you younger scientists)

Hopping and Jumping

Aside from the episode of Dinosaur Train with Cimolestes there are not an awful lot (read: zero/none) of animated or puppeted references to Cimolestes in video, cartoon, or documentary of any kind. There is a nifty little animation that uses the scientific interpretations of the purported movements of the animal. It is a tiny little series of animations of Cimolestes hopping about, which is pretty cool looking honestly. The model is a little more mouse like than I think it should, in my most humble opinion of never having studied this animal in depth at all.

Two Sites Today

There are only two sites needed today. One is About, because it has a nice list of facts and a short paragraph which means it caters to a universal group of readers. The other is the PBS page for Dinosaur Train. The reason this stands alone with the fact page as an assist is because we get a fair number of facts from Dr. Scott Sampson and some inferred behavior discussion. Watching the episode with Cimolestes is also helpful, but there is not a link here for that today; check the PBS site and Netflix for the episode.

Perception of A Tree Rat

©Steve White

Cimolestes was a tree climbing rat sized mammal and it has been drawn as such many times. It has also been drawn to look something like a shrew and even an opossum. Regardless, it is a small mammal with a rodent-like body plan. Adept at clambering up trees and chasing down insects, Cimolestes was an iconic mammal of the Mesozoic. The iconic image of a very small mammal racing around underfoot of dinosaurs is perfectly filled by Cimolestes, however it is usually drawn in the trees, like this image. Its agile little body was slender, but the fluffy images that make it look more like an opossum (not this one) could be said to just be the result of its fur being puffed up. This less furry version looks a little sly, but may be a little more accurate.

Thievery of Bugs

Basal non-placental eutherians are a weird off-shoot of the extant group of eutherians, which are typically known to be placental mammals. One of the best known of those mammals was the Cretaceous scamperer Cimolestes. A North American fossil mammal of the trees the size of a rat, Cimolestes is thought to have chased insects around the arboreal habitats it called home and, probably often, temporary forays into the undergrowth below. Either way, Cimolestes may have been in a transitional place in the fossil history of mammals between marsupials and placental mammals. Additionally, look at how fuzzy and mammaly this animal has finally gotten to be. About time that mammals have started to look like mammals in this jumpy history we have been following lately!

Popularly Not Named

The name of Thrinaxodon is not used in the documentary for which it was used as a model. It appears all over the place though under its own name. Thrinaxodon is included in many other media, including illustrations and toys. The animal appears in popular card games and in the set of collector's cards shown below.

In 1982 South Africa put Thrinaxodon on a stamp also. In a more realistic vein, UT Austin's Digimorph collection is supplemented with a Thrinaxodon skull. My favorite popular culture mention of Thrinaxodon, however, comes from Wizards of the Coast's long standing collectible card game Magic: The Gathering.

Needing Citations

This image desperately needs a citation. Aside from needing a proper credit, it is a great image and very succinctly sums up a lot of the known anatomy of Thrinaxodon. The author/illustrator even included the secondary palate, though it is not visible, which is arguably the most important feature of the entire skeleton in terms of being nearly mammalian. There are other characteristics that show its nearness to the animals that would eventually become fully accredited, card carrying mammals such as the paired occipital condyles, modified jaws with teeth geared toward chewing and the loss of the pineal foramen. There has been speculation that the thorax and abdomen of this animal were separated by a primitive but effective mammalian diaphragm also, which is nearly as significant as the prediction that these animals or their nearest descendants may have been producing milk from mammary glands, a very mammalian trait.

They Wrote A Book

Seriously, they wrote a book on Thrinaxodon. The book is actually only on the cranial anatomy of the small near-mammal, which is slightly more impressive, if that was possible. The book is available as a plain html site from UCMP. That site, in turn, links an even larger work from the University of Texas that was written by Rowe, Carlson, and Bottorff in 1993. I feel like that level of information may be enough for most readers for a day, but there are also published studies on growth patterns and tooth growth and replacement. The writings explain themselves today and are on three of the most interesting topics around Thrinaxodon, which means they require very little further discussion. Enjoy your reading today ladies and gentlemen, there is quite a lot available to you!

Turning to Other Sources

As we have done many times, we have to go to a similar but different species to see anything like what the animal we are discussing in a documentary. Thankfully, the small near-reptile category of lifestyle was well represented despite being an animal other than Thrinaxodon. Labeled simply as "cynodont" in the species list of Walking With Dinosaurs Episode 1, the inspiration for the near-mammal was actually Thrinaxodon though it was not called that in the show. Knowing that, perhaps it is a little bit of misinformation to say that there is not a documentary that has Thrinaxodon in it. The animal is not entirely factual to Thrinaxodon alone however, and incorporates other near-mammals as well. Either way, watching it is beneficial.

Hear and See the Near-Mammal

The fact pages abound for Thrinaxodon. Part of the popularity is in its inherent "cute factor" and the other part lies in its nearness to mammals. Whatever makes it popular, it allows us more of a platform to educate that critical next generation in yet another fossil animal! As noted, there are many sites to look at, including About and the Walking With Wikis which both present the information in a manner that is more accessible for all ages and reading levels. The site Prehistoric Wildlife presents the information as an essay, making it a little less attainable to lower level readers. A video produced by Menteon Learning also lays out this information for us, though does not go into too much depth. This is actually not a bad thing though, as it allows the video to appear concise and accurate in its information, though more information is available than is presented. I am both happy and sad to say that I have another Josep Zacarias link that could certainly be used as a coloring book image as well today. Happy for obvious reasons, but sad because I also found it online without proper attribution, and Mr. Zacarias is a hardworking artist and deserves credit for his work!

Streamlined Digger

Thrinaxodon was built for digging and hiding. The small near-mammal was thin and cylindrical (as far as any tetrapod can be cylindrical anyway) and had a head that was elongated rostrocaudally. This gave it a long skinny body overall and made it look somewhat weasel-like. The major exception to this was the large, but slender, canine-like face with its large canine teeth that appear almost like tusks. The large teeth were not helpful for digging in any specific way, however, the limbs were most built for burrowing. They were short and powerful for digging and burrowing out their homes, but still elongated enough to be suitable for propelling it through the underbrush of small ferns and cycads that populated its world. Given its ability to scurry, probable nocturnal nature, and large teeth, Thrinaxodon was at least omnivorous and most likely fed on insects, lizards, and possibly other near-mammals/mammals and dinosaur nestlings or eggs. Burrowing was an effective way for an animal like this to hide during the day while it ran around at night getting into those dinosaur nests, other near-mammal burrows, and catching nocturnal insets. Efficient as it was then, burrowing is still extremely efficient and useful for small mammals, such as mice and mink, and larger mammals like badgers and otters (River Otters get rather large). It is no wonder that animals like Thrinaxodon would have used burrows for protection. Imagine this small animal shimmying through the hardened soil tunnels!

Breathing and Chewing

Few people stop and marvel at the wonder of being able to chew and breathe at the same time. The ability to breathe while holding food or water in the mouth has evolved a few times in the history of life and each time it has been a remarkable innovation in the line of animals in which it evolved. One of the earliest transitional near-mammals that has been found with the ability to breathe and chew was the Cynodontid Thrinaxodon. Thrinaxodon possessed a primitive secondary palate making this possible. A single species is recognized in the genus: Thrinaxodon liorhinus. This small near-mammal also possessed two small tusk-like teeth and is thought to have lived in riverbank burrows. At under 0.61 meters (2 feet) in length, this small animal would run and hide, use its long whiskers, and come out after dark under the feet of the earliest dinosaurs. These little animals were furry and weasel-like, but they were definitely wonderfully interesting little animals.

©Nobu Tamura

Mighty Moschops

Moschops is a well known unknown beast. That is very confusing, I know. When I have to say that about an animal, what I mean is that no one knows the name of the animal but they have seen the image, somewhere, it happens quite often with prehistoric animals. Moschops has appeared in many popular culture venues, as we have seen. It even featured as a main character in a children's program; that program was a little inaccurate of course. The line drawings of Trainor were even less accurate, overall, though he attempted to keep some of the science accurate. Some old toys still exist, though not many new molds of Moschops have been utilized lately. One of those few new molds has made its way onto YouTube, as toys often do, as a review. Big Time Attic made day #84 in their Dino-A-Day list an entry about Moschops. Their illustration is well done, but kept cartoony, like the Cannons typically tend to do with their artwork (I personally enjoy the child-like aspect of their art).

Calf Face of the Cape

There are four recognized species, though two are considered recognized but doubtful, of Moschops. The heavily ossified skull of Moschops is a defining feature of the animal that may, some speculate, may have been thickly built from birth onward. The speculation states that the fat but short tail adequately counter-balanced the heavy skull of Moschops and that it would have existed with the animal from birth if the head started out heavily ossified. Contrary-wise, if the skull became ossified over time as the animal aged, the counter-balancing tail could have started out miniscule and grown in coordination with the large skull. The skull, in true heavy helmeted-head fashion in the natural world, is very akinetic and appears to lack many visible sutures over much of its surface. The cemented nature of a skull that lacks sutures and kinesis allows for us to infer many behaviors, or at least make some interesting educated guesses, from looking at similar extant taxa with highly "helmeted" akinetic heads (e.g. Bighorn Sheep, American Bison). Due to this, there have been behaviors such as head-butting and display motions inferred in Moschops. The individual pits and bumps on the skull probably aided in the display function of the thick helmeted head of Moschops more than they would have in the physical combat part of intra- or interspecies combat.

Discussing Moschops

In 1936 Frank Byrne of Kansas State University named and described Moschoides which turned out to be a junior synonym of Moschops. It is funny in that Byrne specifically mentions in the first paragraph the large similarity between his new specimen and Moschops. Shortly thereafter, in 1940, Byrne published another article contemplating the evolution of mammal-like reptiles from the Karoo. In that article he again mentioned the similarities between Moschoides and Moschops without tying the two genera together. His discussion of the evolution of the animals is interesting though, despite this discrepancy. On a totally unrelated note, the fighting abilities, which I pondered for a moment or two on either Friday or Saturday, was actually studied and published in 1975 in the first volume of Paleobiology. Herbert Barghusen in Chicago is responsible for reviewing the fighting adaptations of all dinocephalians.

Jim Trainor, Not A Documentarian

Jim Trainor's short film The Moschops is the only short film in his fake "Highlights of the Permian Era" series. The science that exists in it is not the worst science out there, believe it or not, but it is certainly not entirely accurate by any means. The line drawings are typical Jim Trainor (to many that means they are awful). Usually on Mondays I try to only show quality documentaries and resort to not great documentaries where needed. This short film falls into the mostly entertainment category, so attempt to restrain the angry "That's not a documentary or exceptionally educational" comments if you can. The second part of the film, not linked here, gets a little more wayward (i.e. less G rated) with its treatment of the subject matter, hence my reluctance to share the link for that. The problem with a lot of these early mammal-like relatives of modern mammals is that there is not a lot of quality documentary clippings out there. The Walking With Series turns out to be, again, one of the more reliable nearly accurate shows, but it does not exactly discuss the animals we are interested in here today. The non-related Scutosaurus from the series Walking With Monsters probably best shows the posture and a similar gait while Edaphosaurus is probably most similar in diet. Unfortunately, of course, neither is Moschops.

Strange Kid's Shows

In 1983 the British produced a claymation show called Moschops about a Moschops and his friends. Words can hardly describe what that show was, but there are those that swear by it out there on the internet and it may be worth trying to get past the fairly rough models today because it is supposed to be a children's show. A number of episodes exist, but preview for yourself before sharing these potentially confusing and scary looking claymation beasts with your own young ones. The show refers to Moschops as a non-specific dinosaur throughout its run, but nomenclature problems happen in popular culture, so we should probably let this one slide all the way home. Things that are not weird and exceptionally strange also exist. Despite not being a dinosaur, KidsDinos has a typical fact page put up that is useful for relating facts to younger interested parties. They use one of the dopiest looking illustrations of Moschops ever, but that is okay in the long run. About also has a page for Moschops. Thankfully the mammal-like placement and lineage is discussed a little here as well, since it is not discussed in any level of detail on other sites. For those of you that forgot, because it has not been expressly mentioned in the past day or so, we are taking a small hopping path up the mammal family tree starting was back with mammal-like reptiles.

Wrestling Bulldogs

Robert Broom's 1926 reconstruction

In a rather interesting turn of events in paleontology, Moschops has not changed much in stature since the original reconstructions by Robert Broom. Broom's mounted skeleton in the American Museum of Natural History possesses the same posture and pose that modern reconstructions and illustrations bear. The fact that this animal has not been "updated" in terms of posture does not necessarily mean that Moschops has not been studied or better understood in the past 90 or so years. In fact, Moschops has been studied well enough that many genera have been noted to be synonymous with Moschops and have therefore been grouped together. That does not happen without adequate study of the original and subsequent remains. The fact that the posture of Moschops has not been changed in the century since it has been discovered and described means that the posture is actually accurate and accepted by the majority of workers that have been associated with the animal. The posture is actually very much like that of a bulldog (though we know that a bulldog is not a "natural" breed of dog and its posture is somewhat artificial). This bulldog-like position puts the mass of the animal very low and behind the forelimbs, which were massively built. The shoulders are built so strongly in part for traction, as mentioned yesterday, and most likely to aid in intraspecies combat as well as for defensive posturing. It may not have been much of a wrestler, but making itself a much more difficult meal by having a lot of forebody strength would only have made it less susceptible to predators.

Therapsids and Other Early Mammals

©Dmitry Bogdanov

Because of the last month's adventure into the history of the rhinoceros (and the intermediate horse/rhinoceros Megacerops) I have decided that we need to look, as we have done at least once and in a cursory manner, at the other end of mammalian evolution. We will not specifically attempt to trace any single line, however, a fairly comfortable sampling of very early mammals should suffice to examine where some of the familiar forms of today initially took root.

To begin this journey we will look, not in chronological order, but in the order of what I personally find most interesting (unless I get some good suggestions for the future as time goes on). This week, therefore, I have decided that the first ancient mammal to be discussed during December is the genus Moschops. Moschops consists of 4 recognized species, 2 of which are considered questionable. The remains of Moschops species have been recovered from the Karoo Basin of South Africa, long known and famous for its mammalian riches. Moschops was a large, hefty Therapsid Dinocephalian herbivore that possessed a uniquely opened elbow joint that allowed Moschops to move much more like a modern mammal and less like a crawling, sprawling reptile. The skeleton of the animal has led to the inference that Moschops used this newly adapted joint system to move the massive body that the skeleton represents as it fed, seemingly without end, on the low nutritive quality of the cycads and ferns that grew around it. Overall, it is a squat, but gigantic animal. It looks almost uninteresting, but I promise it has many more interesting characteristics to be seen over the week to come.

Well Known Ice Age Beasts

CollectA Megacerops

The fact that Megacerops is so well known that there are a fair number of popular outlets that enhance its popular culture reach. There are toys and references in books, though not in many younger reader books that show up online, and we know it has appeared in documentaries. The only terrible thing about all of this popular culture referencing is that sometimes the lineage of Megacerops still gets confused, lost, or ignored. Many people do not realize that these animals are not exactly and not exactly horses but are related to both; they are more closely related to horses we know. This is very important to note, especially since Kenneth Branagh specifically relates this information to the audience in the Walking With Series. Hopefully we will all remember that in the future since it has been mentioned so much this week.

Ribs and Noses

©Dmitry Bogdanov

Megacerops is, as we have seen, very open faced, skeletally, on the rostral end of the animal. In life, of course, this is not the case at all. That open area is filled with the nasal sinuses and conchae that would have been required by the animal to detect smells sufficiently and to warm the air that it breathed. In colder times of the year this was obviously of great import as cold air in the lungs brings the temperature of the body core down; though being such a large animal to begin with breathing cold air was probably not very dangerous to the temperature of the animal. More interesting topics abound in that nasal area of the skull. We have noted the horns and how they could be used in combat at a pinch, but were they actually of any use? Fossils indicate that damage to the ribs of some larger males could only have been inflicted by other members of the species during ramming contests with the horns. These fractures did not heal properly, if at all, because of the large movements occurring during breathing.

1905, A Big Year

1905 was a big year for Megacerops. Richard S. Lull, of the Massachusetts Agricultural College (presently UMass Amherst), published his description and plates of a fossil he named Megacerops tyleri in the Journal of Geology for the first time. Thirty five years before that the genus Megacerops was originally named and described by Joseph Leidy. Osborn discussed Leidy's description in his 1902 discussion on the Oligocene titanotheres. Either way, Lull's plates depict a skull and forelimb of the animal, far more than Leidy had in his initial descriptions. Prior to this, in the same compile volume, Lull detailed the restoration of the animal. This restoration was effected in clay and features a photograph of the skull, as opposed to the line drawings in the plates of the description. Papers and studies continue to be published, of course. The last well detailed description of a new specimen that I would recommend reading was published in 1995 in the Journal of Paleontology and introduced the new species Megacerops kuwagatarhinus from the White River beds of Montana.

Brontotherium Walking

Walking with Beasts features a battle that is pretty interesting to watch. The confrontation is between Brontotherium (as they are calling it in the show) and Andrewsarchus, an animal known from an enormous solitary skull and thought to be related to pig-like omnivores and potentially whales as well. Regardless, the fight alone can be seen here, with background on Brontotherium. The entire episode is hosted on a different site. However, it may be worth watching to see what other information the BBC presented on these two animals and the other animals that lived at the same time.

Megacerops For Kids

Megacerops, under either that name or Brontotherium is a popular fossil mammal. Multiple sites exist that host lists or paragraphs of facts. About continues to use the more popular Brontotherium on their site while Planet Dinosaur acknowledges the switch, somewhat begrudgingly since they use both names on the title of the page. Brontotheres as a group are addressed on a page dedicated to the fossils of the White River Badlands rather than any single genus in the group. This wider angle approach is good at times and is helpful for viewing all of the animals related to this animal rather than just looking at our target animal. It is always good to look at the bigger picture once in a while.

What Is on Your Nose?

Menodus (junior synonym to Megacerops),
Field Museum, Chicago.

Megacerops has a pair of horn-like protuberances on the rostral end of the skull. In the mounted specimen from the Field Museum, shown here, those protuberances appear smaller than they are often illustrated. The nostrils do not appear, in this specimen, to be completely formed as external foramina, as one would expect. The nares are actually present in the concavity ventral to the the twin horns of the face. The nasal bones appear to extend over the premaxillae between the horns but do not recurve to meet the premaxillae at the chin. As expected in the list of rhinoceros-like traits, the optic foramina are small, relative to the entire skull, and offset laterally so that the animal most likely did not have a great deal of binocular vision. Not having depth perception, we can probably safely surmise that Megacerops was not adept at detecting predators visually. To make up for that deficit we can assume that the powers of smell and hearing may have been more sensitive in Megacerops (there may be more definitive answers that I have not found quite yet). Conversely, mixed herds and even the addition of non-mammalian (i.e. bird) members of the community may have aided in predator awareness, meaning that none of the senses would have had to have been highly adapted toward sensing predators. Either way, the horns on the face of Megacerops are not used for the purpose of combat primarily. As skeletal elements, a broken horn would be tremendously detrimental to the health of the animal. Such a danger would cause the animals to use their horns, both males and females possessed them, as a last resort in protecting themselves. The horns would have served to intimidate as much as the sheer size of the animal itself.

Rhinos or Horses?

Though not a rhinoceros and much more closely related to horses, Megacerops was a very rhinoceros-like relative of horses and is therefore a bit confusing on first glance. Known more popularly as Brontotherium, Megacerops Leidy 1870 was far larger than any modern horse or rhinoceros. Its distinctive look, two large bony protuberances above its nose, make it look slightly more like a rhinoceros, but are significantly different from any known rhinoceros horn as well. The use of these protuberances is well documented, as we shall discuss. This animal is yet another North American Eocene mammal.

Robert Bruce Horsfall, 1913

Star of the Show

After having more than a few handfuls of information written about Paraceratherium and an entire episode of the Walking With Series about one of them, this magnificently large mammal has certainly shown that it has a great deal of popular culture credit. The animal has been shown in video games as well, particularly where it can be modded in, like in Zoo Tycoon 2. It has also shown up in a non-modded form in Jurassic Park Builder as well, to be fair, as a Glacier Park animal. In both games it is seen under the popularized name Indricotherium. There are toys out there, but I would stress caution with buying them under the impression that they are accurate models. Many of the toy images I have seen show extremely muscular giraffe-like animals or emaciated camel-like animals. Considering all of the attention that this animal has received in the last decade, it is very difficult to figure out why the toy models are so ridiculously off base.

What About That Head?

The head of Paraceratherium on the cover of that book yesterday got me thinking. What does the head of a giant rhinoceros ancestor look like exactly? Is it fantastical to have elephantine ears? The simple answer is that we will never really know and artistic license can do whatever it wants to the head. The fact that the illustrator of that book cover went with the head design he did, maybe at the behest of Donald Prothero, only backs up that point. To be fully honest though, that idea is much more innovative than the rhino-on-a-giraffe-neck design that is typically depicted. There is nothing wrong with the usual interpretation because it is usually aligned to the paleontological interpretation of the skull, but with the ears not being preserved on any specimens, to my knowledge, and they could have looked like anything. The open mouth and down-turned lip is sort of mandatory, because of the skull, so the interpretation of that does not often change. Regardless of whether you liked it or not, the new and old interpretations both have merits and you have to admit the idea of a rhinoceros head on a giraffe sized neck is interesting. Seriously though, look at that rhino head on a giraffe neck!

Paraceratherium herd: Elizabeth Rungius Fulda 1923

A Tome of Knowledge

I love original descriptions. That has probably been said many times over. I hate when they are not available for free to anyone and everyone (because it is hard to get articles for some of our younger readers). Despite that happening, there is an absolute mountain of work out there on Paraceratherium. This is due in part to the fact that there are more than a handful of junior synonyms for this animal, meaning that there are a fair number of initial description papers for these now dubious names. There are also new remains still being discovered and described under the currently accepted generic name as well. Rather than listing out hundreds of papers today though, I am encouraging the audience to instead check out, inter-library loan, or even buy a book! This is not because I love mammal paleontology or specifically the work of the author (though anyone into fossil mammals knows the name Prothero), but because anyone that is really interested in Paraceratherium and its family would enjoy and get a lot from Donald Prothero's book Rhinoceros Giants: The Paleobiology of Indricotheres. The first link to the book is a Google powered preview, but this link right here goes to the Indiana University Press. Why would I encourage anyone to buy directly from the publisher instead of shopping around (assuming that purchasing is the intent)? The answer is actually a matter of savings. The publisher is offering the book electronically for $8 less. I love the smell and feel of a new book (almost as much as an old book), but sometimes less is more. You can carry an electronic copy more easily as well. Regardless, I cannot say I have read through this entire book, but the reviews I have seen and the snippets I have previewed point to it being part historical, mostly technical, but very informative overall. As a collector of subject literature (e.g. I get books about birds and dinosaurs whenever I can to compare the information presented regardless of author) I have to say, even if you do not agree with the author, this is a comprehensive work on the animals we have been discussing all week and well worth the investigation on your part! Plus, check out the head on the Paraceratherium on the cover. Pure gold!

See the Models

All of the instances of Paraceratherium are well exhibited in this short video. There is not much to really say about it, but watch this one. The entire episode, if that is more interesting to folks out there, is available here on EntireDocumentaries.

Teach That Memo

Most of the time when a name changes it takes a few years for the internet and sometimes even technical literature to adopt, accept, and print that change. Some of the pages of facts for younger readers today never changed their information and still list animals under their junior synonyms like Indricotherium and Baluchitherium. Any way you pull them apart though, these are all the same animal, Paraceratherium. Enchanted Learning is the biggest offender in that regard, but if one were to go through all of the synonymous names and read the descriptions there would be a wealth of information at their hands. About and the BBC have their naming conventions better put together than Enchanted Learning does however, which is a bit nicer and vastly more convenient. The BBC, of course, uses stills and information from the Walking With Series while About does an even shorter version of what we take all week to do here; integrating technical science and popular sources as much as possible to make the information readable.

Giants and Their Babies

©Roman Uchytel 2010

Knowing that Paraceratherium is the elder synonym of Indricotherium, the name most often associated with this animal, conjures up certain images, mainly those associated with the Walking With Series. The models used for that series are not terribly inaccurate though, to be honest. Some of their models, we have seen in the past, range from not perfect to tolerable with a few outliers in the "oh no, that's not right" category. Regardless, the models for the Paraceratherium/Indricotherium used in the show are exceptionally accurate for a television documentary. The animal is not that difficult to replicate really though. The body is vaguely like that of its cousins, the extant rhinoceroses, but it has an exceptionally long neck in comparison. Its neck is not like that of a giraffe, so calling it a cross between a rhinoceros and a giraffe is not accurate (it has been done). The head is also characteristically rhinoceros-like. As stated yesterday, the best defense for these animals was their size. The babies were already particularly large at birth, but they still required the protection of their much larger mother for a year or so, until a sibling was born, the first calf was weaned, and subsequently shunned b the mother. In much the same way as many large mammals, it is suspected that the mature males were loners that only really socialized for mating purposes, meaning that this illustration is most likely of a mother and child.

He Without Horn

Henry Fairfield Osborn, 1923

Aceratherium is a genus of early rhinoceros that was quite short, without a horn, and fairly unremarkable. Its taller cousin Paraceratherium, however, is remarkably similar but significantly better known. Recognized more readily under the junior synonym Indricotherium. This Eurasian rhinocerotoid and cousin of extant rhinoceroses was one of the largest land mammals of all time. The shoulder of Paraceratherium was approximately 6 m (20 ft) high and its head was capable of reaching another 2 m (6 ft) out from the shoulders. The head itself was approximately 1.3 m (4.3 ft) long. Size, more than anything else, was the primary weapon of this enormous animal. Even their babies are fairly enormous, as we will see at some point during this week.

A Long Horned Devil

I may not have said this this week, but Teleoceras means "Long Horned One". As one of the most common large mammal fossils of North America, Nebraska especially, Teleoceras is also one of the more popular of the "Ice Age" megafauna with the general public. As such, it does appear in a few popular cultural areas including Spore and Zoo Tycoon. The rhinoceros does not appear in many books, strangely enough, and it also does not appear in any documentaries or other video programs. Small rhinoceros toys have been altered by collectors to look like Teleoceras. Unfortunately, that is the extent of the toy presence. Most of the wonderful things about Teleoceras that we have seen this week are exactly how we have seen them; online and in the world of physical fossils only.

Most Common Fossil in Nebraska?

Teleoceras is considered the most common fossil in Nebraska's Ashfall Fossil Beds. Considering that, one would imagine that the information around Teleoceras would be more plentiful. Wikipedia's page is less than a printed page. Regardless, the point today is that the fossil amount in Nebraska is copious enough that the building housing the fossils is called the "Rhino Barn". The state park is home to largely complete specimens, which is even more amazing than simply stating that there are so many remains that there is a building dedicated to housing them. The story of the Ashfall Beds is detailed very well on the park's website. The site is maintained by the University of Nebraska at Lincoln, which may explain why the site is well maintained, in part at least.

Humble Opinions

Scan from the Time-Life book North America (Jay Matternes)

As far as I am concerned there are only two papers that are important today. That is not true, of course. However, for today, the only papers I thought were worth reading are MacFadden's comparison of the Floridian genera Teleoceras and Aphelops and Mead's discussion on sexual dimorphism in Teleoceras. Bruce MacFadden's paper is exhaustive in the most wonderful way. It explores all sorts of levels of the two genera including ecology and niches, diet, and grazing habits. Alfred Mead, meanwhile, explores a single species, Teleoceras major. His in depth study explored the differences in sex, of course, but also in intraspecies habits including herding behaviors and defense mechanisms. Nearly anything that one could want to know, as rudimentary knowledge at least, about Teleoceras is included within these two papers.

Motion Capture-less

Unlike last week's popular Elasmotherium that has charmed its way onto television (and then the internet), Teleoceras has somehow not managed to do so as yet. In fact, it has not managed to make it into any motive media at all. The only mention of it is in a short video of someone showing a Teleoceras molar. It probably is not worth viewing for anyone unless one is that interested in the molars of extinct rhinoceroses (I say this knowing someone that is in fact that interested in that very subject).

Hippo-like for Your Enjoyment

Teleoceras, as mentioned, was an extremely short built rhinoceros. The hippo-like qualities make it much more accessible, or likeable at least, to children, I have found. Regardless, the education component of this post is taken care of in these two PDFs below (Yay for educational material when it is available!). They are actually quite short, but they give students/children something to think about in terms of Teleoceras and extant rhinoceroses rather than just having them color a picture or watch a video. It is a quality endeavor and different versions are available for up to 3rd grade and up to 6th grade. There is one other resource that I like today. The East Tennessee State University Natural History Museum has a neat little page on their site where one can look at the bones of a rhinoceros (labelled Teleoceras sp.). It gives some information about the bones that are being looked at and is generally pretty interesting and worth a few moments of time.

The Weird Eyed Rhino

Heinrich Harder's depictions of ancient wildlife were probably at the top of the world at the time that they were created. To be honest they are still quite beautiful works of art depicting prehistoric animals of all ages and varieties. The Harder Teleoceras is not an exception to this rule. The depictions of Teleoceras over time have certainly changed, but Harder's accurately, perhaps more accurately than many of his other works, portrays the animal in question. The rhinoceros is depicted much like an Indian or Sumatran rhinoceros, but its squat body is what Teleoceras would appear like as well. Remember that Teleoceras has some of the shortest legs of any known rhinoceros, extinct or extant. The only strange aspect of the appearance is the indent behind the eyes. The size of the horn is variable amongst specimens so Harder's depiction is not inaccurate in that respect.

Game of Teleoceras

The family tree of the rhinoceroses has branches occupied by some rather large animals. Elasmotherium we know was a tall and stout animal and the extant members of the family are typically large and stout regardless of how short they might be (Javan and Sumatran rhinoceroses are significantly smaller than their other living cousins). The small members of the family are not limited to these small extant rhinoceroses though. One of the smallest members of the family tree is most famously known from North America (although also known in France), which is not considered, usually, prime rhinoceros country compared to the extant taxa. Often portrayed as a rather portly rhinoceros, members of the genus Teleoceras (consisting of 10 species), were also shorter than their living cousins and ranged the land between around 15.5 and 2.6 million years ago. The holotype species is Teleoceras major Hatcher 1894. Teleoceras possessed a very un-rhinoceros like trait aside from its rather hippopotamus-esque stature, it lacked a prominent horn (let alone a pair of horns) on its snout. Also, the armored skin of this rhinoceros genus is different from that of other rhinoceroses and is portrayed along a line from thick hide alone to thickened armored hide.

Chinese Unicorns

Elasmotherium is repeatedly referred to as a basis for the unicorn myth and moreso in China than in any other place in the world. Due to this there have been entire books dedicated to explaining Chinese unicorn mythology that include hefty chapters on our rhinoceros friend Elasmotherium. These include Jeannie Thomas Parker's book called The Mythic Chinese Unicorn. There are times when the popular culture outlets for Elasmotherium are a little less scientific than even these books about mythology; believe it or not these myth discussions do tend to discuss the actual science behind the life histories of the animals. Regardless, the toys are not always entirely scientifically accurate, however, one of the most accurate is well represented on the internet. The model is from Papo and is reasonably well done, but in this discussion on The Dinosaur Toy Forum the author of the post repainted and enlarged the horn with sculpting material, making for a fairly accurate reconstruction of Elasmotherium. Perhaps the most telling popular culture references are in the multitude of illustrations that have been done in the various styles of illustrators and time periods in which they were drawn. Rather than discussing each style and period of time in which they were drawn, they are presented here in a gallery format:

E. sibiricum ©Dmitry Bogdanov, 2006

E. caucasicum  ©Dmitry Bogdanov, 2009

First published restoration (1878) of E. sibiricum, by Rashevsky

E. sibiricum  ©Philip72, 2002

E. sibiricum  ©Stanton F. Fink, 2011

Rhinoceros Horns

The size of Elasmotherium is greater than that of extant rhinoceroses. The person pictured here is about average height but that really does not tell us much about the height and size of Elasmotherium beyond the fact that it is taller and larger than a person. Then San Diego Zoo keeper Laura Weiner posed near a young Black Rhinoceros and with some Indian Rhinoceroses in the not too distant past that gives us a much better image of the height of an extant rhinoceros next to a human being. Obviously they are large animals and their overall size dwarfs ours, but Elasmotherium would have done a much better job of making us look miniscule when standing next to it. The horn alone was particularly enormous, and fortunately we have materials that allow us to make inferences into what that horn looked like. One such piece of material is the skull at the Natural History Museum of London. This skull has a well preserved base for the horn that allows for well informed extrapolation of the structure. The length of the horn based on that extrapolation may not be entirely correct, but that is a question of math and inferences and topics that I am honestly not well-enough versed in to describe. Either way, these enormous horns would probably have caused Elasmotherium to become endangered just as fast or faster than their extant cousins because of the wealth they would have represented. They would have been pretty awesome to see roaming the plains though, that cannot really be denied.

NHM London specimen User: Ghedoghedo

Elasmotherium on Paper

A multitude of Elasmotherium studies have been conducted since the naming and describing of the initial materials. Limb bones have been described and there have been ecological studies conducted. Geographic populations have been described for areas like China and Kazakhstan. The phylogenetics of the genus have even been studied, probably multiple times to be honest. The original description of the Moscow Mandible and Fischer's inferences about the ancient rhinoceros are lost to the internet, as far as I can tell. That is, of course, rather unfortunate given how important naming and describing articles are to the history of any fossil species and the study of them. Fortunately, however, Zoya Bessudnova recently tackled the history of Fischer in 2013, describing his scientific endeavors in Russia and upholding his fame as the "Russian Cuvier", a term for Fischer I was not familiar with. The most interesting fact about that statement is that Johann (Grigory) Gotthelf Fischer Von Waldheim was German born (1771 in Saxony) and moved to Russia in 1804. Either way, his description of Elasmotherium sibiricum  is not in our hands today, but many other quality papers about the animal are available for reading!

Lack of Evidence

I find it quite disturbing that the movies of Elasmotherium out there are a bit thin. Considering that we are talking about a well known mammal that is a pretty interesting bit of (non)information. One of the only clips that appears widely on the internet that is not game or cartoon related comes from the show Prehistoric Park. The clip is mentioned to be cut from the show, but if it was, that is a terrible idea, because it was very well modeled and looks pretty fantastic.

Not so Much a Dinosaur

Dinosaur Jungle hosts a page describing Elasmotherium that looks fairly whimsical. However, it also looks fairly accurate, so there are not any really large complaints about this either. Even more important, Enchanted Learning has a short synopsis and a coloring sheet. It is not the best, as we typically expect from Enchanted Learning, but it is a rather nice thing to have all in one place.

History in Pictures

Moscow mandible, J. Fischer

The first images released to the public of Elasmotherium have been lost, for the most part. Named in 1808 (Fischer published the account in 1809), that is really a small matter, all things considered. The famed Moscow Mandible was illustrated many times, here by Johann Fischer in the 1809 description. At that time Fischer was the director of the Natural History Museum at Moscow University. The material is quite nice appearing, in this illustration at any rate. Mammal jaws are well known and researched and Fischer's description of this jaw element, which would go on to become the holotype of the genus, was about as typical as one would expect given the material and the time frame of its description. Completely regardless, this jaw and later material began painting a picture of one of the most interesting predecessors of the extant rhinoceroses. Fischer named the ancient mammal based on the teeth that he found in this jaw, making it even more important than if it were just a simple jaw. The name Elasmotherium is derived from the Greek elasmos (layered) and references the tree-ring like layering of enamel on the molars of the jaw. Those molars have large high crowns and are considered hypsodontid. Inferences made from hypsodont teeth include MacFadden (2000)'s statement that "as a general rule, extant herbivores with low-crowned teeth are predominantly browsers and species with high-crowned teeth are predominantly grazers." This is not a new inference, however, as the Russian paleontologist and contemporary/colleague of Darwin, Vladimir Onufryevich Kovalevsky, had proposed a connection between hypsodonty and grazing as early as 1873. The fragment -therium is derived from the Greek therion (beast), simply referencing the fact that this is an animal. The specific epithet of the type, sibiricum, is in reference to the Siberian origin of the jaw.

Heinrich Harder, 1920

Over the decades many things changed surrounding the knowledge of Elasmotherium, including the addition of many more complete, but still fragmentary, specimens of all three species of the genus. The shape of the body and the giant horn took shape. The horn, easily explaining some forms of unicorn legends (though of course these are probably not localized to any one region or type of animal), was enormous and, despite recognizable placement on the skull, was often illustrated in a much more unicorn-like posture. Newer images, we will see during the week, have become much more reminiscent of extant rhinoceroses but older images were significantly horse-like in appearance. This could be partially because of the then emerging knowledge of the relationships between horses and rhinoceroses or it could also be because the body simply appeared that way to earlier illustrators and scientists. Either way, the first full body illustrations were rather interesting because of the horse-rhinoceros cross over of the body shape.

Fischer, J. (1809). "21. Sur L'Elasmotherium et le Trogontothérium". "Memoires de la Société Impériale des Naturalistes de Moscou". Tome II. Moscou: Imprimerie de l'Université Impériale. p. 255. 

MacFadden, Bruce J. 2000. "Origin and evolution of the grazing guild in Cenozoic New World terrestrial mammals". In Sues, Hans-Dieter. Evolution of Herbivory in Terrestrial Vertebrates: Perspectives from the Fossil Record. Cambridge: Cambridge University Press. pp. 223–244