Taller Than an Emu

Bullockornis was obviously a large flightless bird. In the days in which these giant birds ruled the land there were still not very large mammals and the non-avian dinosaurs were long gone. Therefore these largest of flightless birds were the largest animals that were running about terrestrially. The groups of Bullockornis that were living in the Australian Outback were nearly twice the size of the average emu and were capable of chasing down and eating anything that they wanted to eat. This could have included enormous fruit, if there was any; the bill of Bullockornis would have been quite capable of breaking open even the toughest modern coconuts. In the realm of possible carnivorous diet items nothing would be too tough. Bones could be crushed by a bill of that size and strength; the shearing ability would have been fantastic as well. As of now we know almost nothing about the body of the birds, however the best proxy was living at the same time on different continents, so educated guesses can be made with what we assume is great accuracy.

Hell Birds

Bullockornis' original description does not seem to be online anywhere. However, there are a few articles or papers that mention or discuss Bullockornis that are available online. There are only two that I would recommend today though. The first one is is from an online archive of Nature Australia Magazine, published through the Australian Museum. The article discusses the diets of Australia's large terror birds. The second article discusses, in general, the anatomy, diet, and general ecology of the giant birds of Australia. The title puts this much more eloquently, as it states that these "colossal" birds lived during Australia's "Dreamtime".

No Motion

There are no movies at all for Bullockornis. Amazingly, as an enormous bird that had the ability to break quite a few bones with its strong, thick bill, there are no documentaries or stories in general that show this giant bird in action. We could, of course, watch the various "Terror Birds" that have featured in documentaries, fake nature shows, and even somewhat serious television dramas. Sometimes that is just all we can do.

Ducks for Kids

Farm ducks tend to make tolerable pets for some. This duck would not. Bullockornis does, though, have many places that we can learn facts about the genus and share them with kids. It would probably also make for a really interesting stuffed animal, but I haven't looked to see if such exists yet. However, I have seen that About, Prehistoric Wildlife, and even a short blurb from the government of New Zealand (featuring a painting of a different genus) can be found online.

Demon-Duck of Doom

(C) Peter F. Murray

Maybe the best part of the name Bullockornis is that it is not the name by which the animal is colloquially known. The giant, heavy bill of the Miocene bird in addition to the proposed phylogenetic sisters of the bird (ducks and geese) lent their influences to the nickname Demon-Duck of Doom. It is with that nickname and the known fossil material in mind that many of the illustrations of the enormous flightless bird have been penned. The most coy looking version in the world once graced the pages of National Geographic while showing off its enormously powerful beak. The lower portion of its body was loosely based off of the other known Terror Birds of the southern hemisphere. There is probably a little bit of the Eurasian Terror Birds mixed in there; however, this Bullockornis is most definitely the most smug extinct animal to have ever been illustrated in the pages of National Geographic.

Could Not Resist

Perhaps not the most popular of the giant flightless birds of the Miocene, but certainly an enormous bird well worthy of being considered a "terror bird" (a name generally associated with the South American birds), Bullockornis planei (Rich 1979), was an Australian monster. The name refers to Bullock Creek, where the fossil skull was discovered. The material consists of a rather large head, thereby making the name, which translates to "Ox Bird", doubly meaningful in addition to the locality reference. As someone interested in avian crania, this is a fantastic fossil find with tons of potential. Also, look at the beak on this bird? It is awesome, in the truest sense of the word.

(c) Gord Webster

Popular Frogs

Partly due to its size Beelzebufo is a highly popular amphibian. The majority of its popular impact, beyond television and pure knowledge of its existence has actually been through the game ARK. Dinosaur Revolution and Dinosaur Train are the prime television conveyors of knowledge about Beelzebufo. It did leave behind a rather fantastic skeleton either way.

Portmanteaus are Fun

Previously I mentioned the origins of the name Beelzebufo ampinga, but not why the portmanteau was chosen. The specific epithet of the Malagasy word "shield" is reflected in the shape of the head of the frog and also in the fact that there is some evidence that the dermatocranium of Beelzebufo may have had some rough scales or dermal scutes protecting the top of its body and head. The rugosities at the very least point to a dermally protected head, but we can hypothesize that this scuted armor potentially extended caudally from the head as well. This may be the least drastic hypothesis concerning this frog ever. The devilish part of the frog's name comes from its hypothesized relationship to the extant genus Ceratophrys (and the family encompassing the genus, Ceratophryidae) which includes frogs with supraorbital horns accentuating the apex of the head. This explains why many of the illustrations of Beelzebufo include supraorbital horns. The relationships, though, are more important in that Beelzebufo has been used as evidence that frogs in India, Madagascar, Africa, and South America are all related, partially through Beelzebufo and that, therefore, their ancestors all once inhabited the super continent Gondwanaland which must have split apart, isolating Beelzebufo and the Ceratophrys' common ancestor sometime during the late Cretaceous. That common ancestor then differentiated over time into the ancestor of extant South American Ceratophrys frogs and Beelzebufo. I think what we all really want is a time machine so we can see these changes because frogs, often forgotten by the populace, have a rich and intriguing history that is not terribly well understood, though we can say that about even the best understood lineages.

At the Bottom of the Paper Pond

Papers that come out for a new species beyond description usually fall into a few categories. There are phylogenetic and familial papers. These can be interesting reads as they take the descriptions one step further and discuss how those descriptions relate to other fossil and sometimes living taxa. There are "new material" papers that add to the description of the original material and sometimes add to our knowledge of the skeletal anatomy (rarely but wonderfully they can also add to our knowledge of the soft anatomy) of animals. The description of new material of Beelzebufo is significant because amphibians have a sketchy fossilization ability as is. The original material of Beelzebufo is actually nicely preserved and somewhat substantial, but additional material is never a bad thing. Strangely, searching Google Scholar, the hits on Beelzebufo very quickly become papers with snippets about the frog in Spanish, Italian, and what I think may be Norwegian (it is certainly Germanic). However, the three papers above are more than enough for a day's reading on a giant frog; if there is such a thing. Reading about giant frogs is actually quite interesting and fun.

Beelzebufo the Terror

The tiny terror toad tormented tribes of toy theropods throughout the Cretaceous. All alliteration aside, Beelzebufo was an impressive ambush amphibian and quite capable of terrorizing the smaller dinosaur hatchlings, lizards, mammals, and "lower vertebrates" that lived alongside it. Invertebrates were most definitely also on the menu. However, in this popular clip from Dinosaur Revolution, the smallest theropods or lizards are the main course. Strangely, despite all hypotheses to the contrary, these Beelzebufo use their tongues to attack their prey items; a mode of attack thought to be the mainstay of other frogs, and not the gulping of the gigantic mouth of Beelzebufo.

Strange Occurrences

Flipping through the stations as I type PBS is airing the episode of Dinosaur Train featuring Beelzebufo. Netflix still has a lot of Dinosaur Train episodes, so this may be able to view on demand. I would recommend watching it, not only because it is available, but because it is geared toward a younger audience, which is great. Either that or you can turn on PBS in the next five minutes. The best part about the character on the show is the fact that they made him sound a lot like a mob boss. National Geographic has a page on the giant frog that is a bit more intermediate which is not too difficult to read. Prehistoric Wildlife is the only other substantial page that we will share today.

It Eats Everything

Most likely because Beelzebufo is so enormous, again, for a frog, most of the illustrations that exist of the fossilized amphibian are depictions of the animal eating some kind of large lizard or a very small dinosaur. Many of the illustrations also make the frog appear very similar to a horned frog or a pacman frog (both general common names belong within the genus Ceratophrys). This is probably an echo of living species that artists have to draw from. The skeletal elements are also somewhat similar to the large frogs that are extant and, a good portion of which, roam South America now. Regardless, at 40 cm (16 in) Beelzebufo was quite capable of eating small and large lizards, mammals, and even hatchling dinosaurs as they scampered through the bushes. Anatomically this frog was a frog, so the snatching and eating of its prey was probably, as in many of the enormous extant frogs, pursued through ambush and large gulping instead of using a sticky tongue like some other extant frogs.

Terrifying Frogs

All of the thoughts and discussion on amphibians the past week made me really want to look up some of the other groups of amphibious animals that are represented in the fossil record. As much as we all claim that the fossil record is awful and does not do much for us in the way of preserving enough life, there is a lot of quality frog data in the fossil record. One of the latest and largest finds is a frog that has been named Beelzebufo ampinga. The genus name is a combination of Beelzebub and bufo, the "Lord of the Flies" from Semitic culture (not meaning specifically Hebrew culture but the collective cultures of southwestern Asia around the time that Hebrew culture was "born") and the Latin word for toad, respectively. The specific epithet, ampinga, comes from the Malagasy word for shield. The enormous frog weighed in at 40 cm (16 in) and 4 kg (9 lb), larger than any living frog. The fossils came to light in 1993, but were not described until 2007, and as such the giant frog is new to popular science. Even if it was not, its enormous size, for a frog, make this fossilized amphibian popular and intriguing. The image and caption below is from the following reference:

Evans S, Groenke J, Jones M, Turner A, Krause D (2014). "New Material of Beelzebufo, a Hyperossified Frog (Amphibia: Anura) from the Late Cretaceous of Madagascar". PLOS ONE. DOI:10.1371/journal.pone.0087236. PMID 24489877. PMC: 3905036.

Three-dimensional digital reconstruction of skeleton of Beelzebufo ampinga highlighting sources of material for reconstruction. A, dorsal view; and B, right lateral view (with left limbs removed for visual clarity). Beelzebufo specimens used in model in dark blue. Light grey cranial and vertebral materials inferred from known morphology of Beelzebufo specimens, primarily through mirror-imaging. Dark grey jaws and postcranial elements modelled on large female specimen of Ceratophrys aurita (LACM 163430)

Where You Know Him From

Given the fact that Diplocaulus does not appear in many documentaries or other popular outlets, it seems somewhat odd that Diplocaulus is a popular animal. In part, the shape of its head has helped quite a bit, considering that it is extremely unique in not only the animal world but the salamander and amphibian world much more specifically. We have seen that Diplocaulus appears in many video games and its odd shape has been described many different times by many different people. Multiple descriptions may be part of the reason so many people know about the animal. It is probably, though, the fact that the fossils are relatively common, overall. We can all admit that it is the wonderfully weird shape of the head. The most interesting thing about the head of Diplocaulus is that it may have been used, in part, as a hydrofoil. That hypothesis is somewhat less popularly publicized, but whenever it does surface it is rather popular. In fact, those tabular horns, as they are more properly known anatomically, were widespread throughout the order Nectridea.

Devil Horns

Diplocaulus means "Double caul". A caul is a derivative of the word cowl, and, without surprise, the name refers to the doubled cowl-like horns that protrude caudally from the head of Diplocaulus. The majority of fossils of Diplocaulus include little postcranial material, as is typical with a great deal of fossil materials that have been collected. Considering the lifestyle of the amphibian it is amazing we do not have more well preserved slab specimens. This sort of specimen, with its typical amphibian (salamander-like) body shape, is a spectacular specimen, though not all original material.Probably the best maps of the anatomy of Diplocaulus are actually the skull maps that are based, typically, off of the original descriptions by Cope, Olson, and Williston. See that map of cranial bones below. Obviously the amount of unfused bones, in comparison to a mammal, is a lot larger and impressive.

Sources Everywhere

Diplocaulus has made it into books that discuss its relationship to sister taxa and structures, though 1917 is one of the last prominent examples of this. Many examples of the fossils exist. Enough, in fact, that growth series of the lepospondyl have been stastically analysed and the number of stages in that growth series have been hypothesized. My favorite, of course, is the S.W. Williston description of the skull and extremeties of Diplocaulus. In true Williston fashion the articled is back-ended by quality hand drawings of the skull and parts of the limbs from multiple angles. He even includes a plate labelled with the bones, which is always nice.

Unhappy with Documentaries

Possibly one of the saddest days is when we have an animal, of any caliber, that would make a great addition to any documentary but is featured in none. Diplocaulus, for all of its uniqueness, is one such genus of animals. The amphibians are curious and intriguing characters, but have never made a key appearance in a high profile documentary that has then made a lasting impression on the internet or television. I cannot think off the top of my head, of an appearance that was in a popular program either. However, there are videos on the internet in which Diplocaulus of one species or another features. Many of these are video games, as the animals appear to have made an impact, somehow, in the Jurassic Park world. They are popular characters in fighting versions of that world in particular. The best videos though, are probably the ones that claim that Diplocaulus is still alive and roaming the wild. This video is one in that vein:

Listed Amphibia

Rather than wax poetic about the number of links, which is many, here are a few for today:
KidsDigDinos
Dinosaur Jungle
Enchanted Learning
About
KidsDinos
There is also a fun little sculpture time lapse, like the one I shared with Protostega a couple of weeks back. It is very nifty and enjoyable. Part of the reason this timelapse is more interesting, though, is that the sculpture is not done by hand, but in the free to use 3D software Blender.

Alternative Heads

Display at the University of Michigan

The Diplocaulus magnicornis on display at the Natural History Museum at the University of Michigan is a special duck, as my mother sometimes says. The fossil is as about as normal as any other Diplocaulus fossil that has ever been found. The head and vertebral column were well preserved while the shoulder girdle, pelvic girdle, ribs, and all limb elements have been lost in this specimen. The most interesting thing about the specimen is actually the hypothetical reconstruction that accompanies it within its display. The flattened head, in this hypothetical recreation, supports a petagium that extends back over the body, attaching to the caudal portion of the pelvic girdle. This petagium makes the robust little salamander look much more like a squid than an amphibian. If this morphology was even a partial reality, a muscular portion of the attachment might have made Diplocaulus almost part skate or ray in the way in which it swam. A salamander-like animal undulating through the water would be interesting, but it probably was not very likely.

Tribulations of An Aerodynamic Head

©Dmitry Bogdanov

The salamander-like body of the genus Diplocaulus (with two accepted species: D. salamandroides Cope, 1877(type) and D. magnicornis Cope, 1882)
was meant for walking about on the bottom of the aquatic landscapes that the lepospondyl amphibians in the Order Nectridea called home. Their heads, however, were built in a very interesting manner. They were, quite obviously, the most interesting things about these Permian animals of North America. Many hypotheses have been formulated about the way that Diplocaulus swam, ate, and foraged or hunted in the rivers, lakes, and swamps that they inhabited. Habitats of Diplocaulus have been narrowed down mainly to these simplified definitions of freshwater systems. Regardless of these definitions, as the largest known lepospondyl it was restricted to the deepest and widest of these freshwater habitats. Supposed remains from Morocco have made this enormous salamander an intercontinental, in the modern global view, fossil that may be able to provide further evidence of continental drift. That is, of course, if those supposed remains belong to the genus Diplocaulus or one of the known and accepted species.

Popular Near Fish

Haikouichthys has not been ignored by the public. Part of the reason behind that is that it played such an important role in evolution both on television and in the actual process. These animals were not as charismatic and happy looking as they appeared on television, as far as we know, but they are well known and studied at this point. Although, the happy little eyes that  we can see in this image are very interesting and happy looking.

Heads and Tails

Nobu Tamura

The fact that basal chordates like Haikouichthys and Myllokunmigia (a contemporary of our animal this week) had crania and tails that are visibly differentiated and identifiable by anyone, even those with very little anatomical experience. The head is not extremely noticeable, as with a giraffe, for example, but in most illustrations and fossil slabs the small animal can be seen to have a head-like end with small eyes on the end of that small head. To be certain, Haikouichthys was a small animal with a minuscule head, but it is certainly a head. The notochord is not as apparent in the illustrations but is apparent in many of the known fossils. The cord is simple and as basic as a noticeable spinal cord could possibly be while still being called a notochord. The first spinal cord was still a ways off, evolutionarily speaking, but this simple hollow dorsal nerve tube, one of the first we know of for certain, was an enormous step on the way to every living vertebrate, not to mention countless fossil species, that we see and interact with every single day.

Notochords and Potential Fish

Haikouichthys is probably one of the most discussed almost fish in the entire fossil record. It is probably one of the only almost fish in the fossil record that has enough specimens available to researchers to be worth the time and effort that is spent on them anyway. Thankfully, though, they are worth that effort and many a quality article has been written about Haikouichthys. These include descriptions of the basal cranium and notochord and the fin structure of the tiny ocean-dwellers. These two articles are pretty heavy and, therefore, are all I am going to share for the day. Enjoy the articles and the rest of the night.

Short and Sweet

The shortest video ever is what Haikouichthys boils down to in terms of documentaries and video. That may be a little hyperbolic actually, but it does indeed have a very short segment of the BBC special that followed Walking with Dinosaurs, Walking with Monsters. Monsters told the tale of life prior to and up to the dawn of the dinosaurs. The segment featuring Haikouichthys is pertinent to the history of life as the development of crania and notochords are both key elements of that history.

Dodging the Scary Seas

There are a lot of links in the internet ocean in which Haikouichthys is floundering. There are tons of bad links dodging in and out o the good links and therefore navigating the sites is kind of a task on its own. The problem, simply and quickly put, is that animals like Haikouichthys are often the center of evolution/anti-evolution debates and arguments. There are plenty of good sites though, such as About, Palaeos, the Walking With Wiki, and Prehistoric Wildlife. The Prehistoric Wildlife site actually has a good line drawing that can be used as a coloring sheet as the precarious position of Haikouichthys is discussed with others. The site also has a size comparison image that, if one did not know the size of Haikouichthys before, is fairly fantastic.

Description and Labels

Paleontology has a great habit of pairing quality photographs with highly informative labels of line drawings. This happens, of course, in the best papers and less often in papers that are not as high in quality. In fact, it is the highly detailed photographs in conjunction with highly detailed text description that often make the best papers when it comes to discussion of anatomy and characteristics. We are fortunate with descriptions of Haikouichthys because many of them include detailed description, photos, and labels on line drawings. Sometimes the fossil can be adequately labelled on its own, though. Today we have an adapted image of a number of views of fossils that show different aspects of Haikouichthys. In that image we can see different views of the eyes, fins, and rostral end of the animal. The most interesting aspects of the fossil are the shape of the dorsal fin and the tail fin. Also of interest are the clearly visible myomeres, which are undifferentiated muscle bundles that are, effectively, the most primitive form of musculature an animal can possess. We do see myomeres in extant agnathan cyclostomes such as hagfish and lampreys. This level of organization is, strangely it seems, still evident therefore. That allows us to have similar, but not identical, locomotive systems in animals that are still living in aquatic environments. That type of diversity is actually quite fantastic.

A Different Kind of Fish

I wanted very much to discuss the basal skate/ray Heliobatis this week, unfortunately the information that exists for the little fossils is not very good. The fossils themselves are plentiful and beautiful, so we may venture back to the topic, but this week we are going to discuss what is, on television, considered one of the more important fish in the history of evolution. All fish are actually very important to our history. The reasons that all fish are important is that a lot of what life has become on this planet can be traced back to fish, including fish without mouths. How does a fish not have a mouth one might ask. This is certainly one of the more interesting questions of science. As with any evolutionary question, the line of evidence and the existence or absence of traits is sometimes shocking to us as we look at living animals to try to understand fossil animals. Animals have, through their history, gone through forms without heads to forms with heads, then they gained distinct mouths, and eventually teeth. That very cursory history of the development of jaws and teeth is lacking, but the general path is well known. The small fish Haikouichthys fits into the very early part of that development. Cladistics argues against Haikouichthys placement on the trees as one of the earliest fishes, but does agree that it could be either a basal craniate, animals having clearly defined heads and skulls, or a basal chordate, animals with spinal cords, often referred to as notochords in the primitive conditions. Since we have spinal cords (the more "advanced" version of the dorsal nerve cord or notochord) and clearly defined skulls, somewhere along the line we are actually related to this small jawless fish.

What is a jawless fish? How can a fish be jawless? Since we are accustomed to extant animals having jaws (for the most part), it can be difficult to entertain the idea of a fish without a recognizable skeletal jaw. Appearing somewhat similar to a hagfish, Haikouichthys had an opening where a skeletal jaw could have been, if it had existed, but was instead little more than an open vent through which the tiny Cambrian fish could filter food. Lacking jaws even made sucking in food impossible for agnathan ("no jaws" from the Greek) fish. As we shall see, this kind of life was difficult, but even more things in the Cambrian oceans made life more difficult for this small fish.

King Turtle

Aside from Archelon, Protostega is probably the most well known fossil turtle in existence. The fact that it has been known since the late 19th Century partially is responsible for the wide circulation of knowledge connected to its existence, but it is also a byproduct of the wholeness of many of the specimens that have been collected. This is thanks in part to the marine environment in which the turtles lived, but many of these remains also probably represent turtles that, unfortunately for the living organism, were killed and buried quickly if not simultaneously. Marine animals tend to bloat with gases and float near the surface, allowing scavengers to pick apart the carcasses before they settle back to the ocean floor. While this certainly happened to some of the giant sea turtles it may not be the case for all of the extremely well preserved fossils. The fame of the turtle in our culture is partly owing to the museum displays that have been erected to show the fossils as well as quality sculpted versions of the turtle, but the other major factor in the popularity of this turtle is most definitely the fact that it is an enormous sea turtle. Sea turtles have always been much beloved by many people, and fossil turtles are no exception. Whenever we see a sea turtle swimming in a local aquarium or the ocean we can imagine a larger version of it swimming about, which is basically what Protostega was.

Sizes of Gigantic Turtles

©Dan Varner (via Mike Everhart's Oceans of Kansas)

Unfortunately some of the greatest paleoartists of all time are in the past (though many still thrive and survive now). Dan Varner is certainly amongst those numbers that have passed on but he was relatively unknown to general audiences. The main reason is that his art was concerned with the Western Interior Seaway and the creatures that were living there in its heyday. Thankfully, though, he illustrated wonderful and accurate (as accurate as possible) animals that we still discuss to this day, like Protostega. This illustration, for example, shows the sheer size of an adult Protostega in comparison to an adult Hesperornis. To those without the knowledge of Hesperornis size, an adult bird in the genus would reach approximately 5ft from beak to tail. The fact that the bird barely reaches from one side of the turtle's shell to the other side leaves us with an interesting view of the enormity of the turtle. The carapace is obviously longer than it was wide and, adding on the flippers, the total width of the turtle was most likely in excess of 10 ft in even the smallest adult individuals. As we stated earlier, adult specimens of Protostega gigas were almost entirely safe from the apex predators of the ocean. The babies were probably in danger from the laying of the egg until it reached the adult size however, as with many extant marine turtles.

Turtle Descriptions

Turtle phylogenies are ludicrous things. The trees with turtles well represented are biased one way or another; toward archosaurs or closer to lizards, generally. In 1898 that debate was just starting to become highly published, in relation to the species Protostega gigas. O.P. Hay was one of the first to describe the relationships between turtles like Protostega and other turtles like Dermochelys. This was after, of course, the initial description of the original fossil material by E.D. Cope in 1871. Descriptions abounded early in the knowledge of the existence of Protostega beyond these. Charles Sternberg even described new fossil materials of Protostega gigas at the turn of the century, or shortly thereafter, in 1903.