Holidays and the New Year

Good afternoon all. As promised, in the new year we will see a new name and some new branding to this site. We will not forget dinosaurs entirely, but we will widen our scope to more and more fossil animals from many different families. This week, however, there is going to be a short hiatus and next week, rather than a specific animal, we will look at the new images and banners (provided I put something together in time) and during the first week of the new year I will have a bit of a surprise as far as topics, but that can wait. For now, though, here are some old holiday images I have posted in the past. Regardless of what you celebrate, happy holidays and enjoy your week off school or work or whatever!

Few Images

Han, et al. 2015

There is a lack of images for Hualianceratops for one main reason: there is very little skeletal material known and that material is almost all in the middle portion of the skull. Any ornamentation or odd body shapes are unknown to us with the material we possess at this time. Therefore, instead of showing a really imaginative or well done image that we have not seen this week. Instead, you will find an image from the descriptive paper summarizing the placement of the bones within the living animal's skull. This reconstruction of the skull labels the known material of the skull also; quadrate (q), postorbital (po), jugal (j), maxilla (ma), dentary (d), and predentary (pd). The scale bar of 5 cm shows us how small this animal was as well. The entire head was not much longer than about 15 cm, which is fairly small for what we think of dinosaurs. However, there were many small dinosaurs about this size and, for many lineages, starting off small was quite normal before we see the enormous hulking dinosaurs of later generations, in this case giants like Triceratops.

Short History Lessons

Many partial remains do not generally introduce a large amount of independent study in the annals of publication. The reason, of course, is that partial remains do not contain a lot of material to be described or to guide inferences into form or function. The material that is known for Hualianceratops, for example, is a minimalistic (not in a derogatory way) collection of cranial elements from a single individual that have not been studied much beyond the scope of the initial description and phylogenetic comparisons (newly published) made in other papers by some of the original authors. The original description, remember, was the work of a collaboration between paleontologists associated with the Institute for Vertebrate Paleontology and Paleoanthropology (IVPP) and George Washington University published through The Public Library of Science as Han, et al. 2015. As stated earlier, there are few studies of this animal outside of these two studies. One study that has specifically mentioned and discussed Hualianceratops in a different context, though, is Maiorino, et al. 2017. This paper discusses biomechanical function of ceratopsian dinosaurs during feeding. What that means, in layman's terms, is that the paper describes how ceratopsian dinosaurs eat their food. The paper also addresses differences in the form of the ceratopsian skull across species.

Longer Videos Today

Rather than go into all kinds of news sources and random short videos, today I will quickly share a longer video that discusses the (then) newly described dinosaur Hualianceratops. Enjoy the news and facts presented in this short video and share it around.

The Oldest Ceratopsian

In the Late Jurassic the foundations of the anatomy of a group known as the Marginocephalians was being laid in the ancestors of these animals. One of the oldest recognized ceratopsians, one of two prominent groups that emerged in the Marginocephalia, was recently (2015) described from remains discovered in China. Institute of Vertebrate Paleontology and Paleonathropology (IVPP) V18641, better known as Hualianceratops wucaiwanensis, was recovered in Western China, specifically in the Xinjiang Province in 2002. The expedition that uncovered the remains was led by the IVPP in conjunction with George Washington University of Washington D. C. The description was penned by Han Fenglu and Xu Xing associated with IVPP and Catherine A. Forster and James M. Clark of George Washington. They described a ceratopsian dinosaur the size of a medium sized dog with a name meaning "Ornamental face" (Hua and lian) with ceratops still meaning "Horned Face". As we will see this week, the remains are heavily based on cranial material, so a name that describes the face twice is actually extremely appropriate.

Before and After Images

Hesperonychus, as we mentioned, appeared in a single movie and has not appeared in many other venues at all. There are some illustrations online and in dinosaur encyclopedias and other books. Many of the artistic impressions that exist of Hesperonychus are very similar in many ways. One of those interpretations was used to create the models for the movie screen Hesperonychus and the before and after images of this concept are similar, yet strikingly different from one another. The initial illustrations for the movie were a collaborative effort, as were the 3D final models, between artists. One of them, Philip Whiteley, has hosted the image seen here on his website showing the concept work and the final product. Notice the dimensional differences and the fluffiness that was enhanced in the final model. The feathers were apparent in the concept as well, and appeared fluffy, but seeing them retained and look alive in the movie was fairly fantastic.

©20th Century Fox and Animal Logic

Descriptions of Small Dinosaurs

Hesperonychus has not made much of a buzz since it was initially described. That paper, shared on Saturday initially but here again today (Longrich and Currie, 2009), includes the expected high detail photographs of the type material, but it also includes museum material hypothesized, but not necessarily known, to belong to Hesperonychus. The conclusions that caused this material, isolated pedal phalanges, to be referred to this species, was tentative at the time and has been based on characteristics of size and shape that place the toes in the same family and, because of the size, attribute them to an adult animal approximately the size of the estimated adult size of Hesperonychus. These phalanges are described and compared to the phalanges of other taxa in order to justify their reference to this species of dromaeosaur. Key elements of the pelvic girdle that mark the animal as an adult stand as diagnostic characters that separate Hesperonychus from other genera and the possibility of the small dromaeosaur belonging to another genera as a subadult or even as a juvenile. The characteristic growth described here is fusion of the pubes and ilia, a morphology seen in "somatically mature" animals, as Longrich and Currie state. A phylogenetic discussion is presented as well in an attempt to ascertain the precise clade to which Hesperonychus belongs and its position in both this specific and the larger dromaesaurine clade as well.

Hesperonychus On-Screen

Hesperonychus has made one appearance on a main screen since its description. A clip of the Walking with Dinosaurs movie, shown here, displays their introduction early in the film. If you want to see more of this small carnivore in the movie, I would recommend watching it. The film was produced by the BBC, in part, and some of the same people behind the BBC's television program of the same name. Additionally, this movie is one of the very few dinosaur movies that aims at more realistic animals, including feathering on their animals, among other things.

Facts About the Western Claw

Hesperonychus facts pages are somewhat rare on the internet. Rather than going through the few that show up individually, here are a couple that are worth reading today:
DinosaurFacts
BBC

There are no fact videos hosted anywhere online. There is a tribute video on YouTube that can be viewed. A lot of the images used in this video, though, are either generic or are not the dinosaur Hesperonychus; the latter are not identifiable enough to recognize at least. The images loop throughout the video. Enjoy!

The Western Claw

Discovered in Alberta's Dinosaur Park Formation's Cretaceous strata representing the Campanian stage of 76.5 MYA, Hesperonychus elizabethae is a small dromaeosaur known from a pelvic girdle discovered by Dr. Elizabeth Nicholls of the Royal Tyrrell Museum in 1982. The skeletal element remained in storage until 2009 when it was described by Longrich and Currie (A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America). A number of claws that appear to be allometrically and anatomically related to Hesperonychus, but these have not been described or officially attributed to the small dinosaur. Measuring in at an estimated length of 1 m (3.3 ft) and weight of 1.9 kg (2.2 lbs), Hesperonychus appears to have been a fully adult but one of the smallest carnivores of North America. Despite the small amount of material, the description relied on comparison between this pelvis and the pelves of other dromaeosaurs.

©Nobu Tamura CC BY 3.0

Drawings of Suzhousaurus

©Masato Hattori

Suzhousaurus is a fairly typical looking therizinosaurid theropod dinosaur. Because Suzhousaurus looks like any other therizinosaur in almost every interpretation that exists, this rather large theropod, estimated to measure approximately 6 m (19.6 ft) and weigh up to 1.3 tonnes, is not the most exciting dinosaur illustration subject. However, the least static of these paleoillustrations is Masato Hattori's low placed, high angled interpretation of Suzhousaurus standing behind short grassy vegetation. This interpretation has a shorter coating of feathers than what is generally depicted on therizinosaurs, but the feathering appears soft and downy; Suzhousaurus may not have been as fuzzy and warm as it appears here, but I do not think that should bother us. It is feasible that Suzhousaurus was more likely to be found in woodlands or at the edges of forests than in a plainscape, as this appears to be, but this view may not mean anything in terms of where the Suzhousaurus is supposed to have been living at the point in time that this illustration may be depicting.

Papers and Anatomy

In 2007 Suzhousaurus was officially described and announced to the world as a "Large Therizinosauroid" of Northwestern China. The description available online through Acta Geologica Sinica (the English edition for those that cannot read Chinese characters). The description includes detailed color photographs of the skeletal remains as well as maps of the region from where it had been recovered. Therizinosaur specimens and their stratigraphic ages are also cataloged in the description paper. The phylogeny of the therizinosaurs is discussed briefly, but only enough to describe where the new dinosaur Suzhousaurus fit into the family tree. The description of a newer specimen in 2008 by Li, et al. contains more descriptive information about individual skeletal elements of this new Suzhousaurus megatherioides specimen that was discovered in the same general locality as the type specimen (Gansu Province). The diagnosis of some of the material is revisited and redescribed as well as the new material. However, the heart of this description is in the new material and stretches in great detail for a number of pages. Comparisons to other therizinosaurs round out the discussion, but the overall phylogeny of therizinosaurs is not broached in this paper.

A Presentation for All

This video has an oddly hushed voice reading the facts and offering interpretation, but the strange huskiness of the presenter is a little odd in spots. Despite that, of course, there are a lot of important facts about the therizinosaur Suzhousaurus that are contained within the video. Clearly it was a wide-cast net for a project, but it is a lot of information that is very important for better understanding of the dinosaur. Fact pages on the internet are found sparingly, but there is a Live Science popular science article from around the initial time of publication that helped to announce and disseminate some of the facts about Suzhousaurus.

Chinese Therizinosaurs

©Michael B. H. under GNU Free Documentation License

Suzhousaurus megatherioides (Li et al., 2007) is an Early Cretaceous therizinosaur discovered in Gansu, China, a region in the northwestern portion of the country. The name refers to Suzhou, the ancient name for the Jiuquan area of Gansu where the fossil was discovered. Interestingly enough this region is the northwestern region of this northwestern province. The specific epithet references the similarity of the postcranial fossil to Megatherium, the genus of Giant Ground Sloths. A significant portion of the skeleton has been described. There is an interesting phylogenetic confusion about Suzhousaurus and the lost "Nanshiungosaurus" type specimen; a thereizinosaur genus described with two species (N. brevispinus Dong, 1979 (type) and N. bohlini Dong and You, 1997). We will attempt to look more deeply into this later this week. Suzhousaurus currently sits in a basal position within the therizinosauridae and will therefore tell us a lot this week about what defines therizinosaurs in relation to other theropods. We will find, additionally, what makes the diet of both Suzhousaurus and other therizinosaurs different from other theropods.

Rodents of North America

Reconstruction of the skull of
H. andersontau from Korth 1992

The final animal this week is a small rodent known as Hitonkala contains two species: H. andersontau Macdonald 1963 and H. macdonaldtau Korth 1992. The meaning of the name is very straightforward for this animal. Though some of the other species we saw this week had names that were multi-part translations, Hitonkala simply means "mouse" in Sioux. Hitonkala andersontau is known from isolated teeth and four skulls and H. macdonaldtau, however, is known from a fairly large collection of skulls, teeth, and mandibles. The descriptions of both Macdonald and Kroth detail the teeth in minute detail, but there is not as much description space dedicated to the skulls or any of the postcranial material (I have not found any described at all unfortunately).

The Anthracotheriidae

Type specimen from Macdonald 1956

Anthracotheriidae were hippo-like ungulates found globally that are related to the hippopatomuses and whales. The earliest specimens originated in Asia and some made it to North America, but they are best known and were most prolific in Africa and Europe. One of the species that has been found in North America has a Sioux name, again bestowed upon it by J. R. Macdonald: Kukusepasutanka schultzi Macdonald 1956. The name means "Big hog nose" and Macdonald, for this genus, does not include a pronunciation guide; though he does thank Mrs. Lulu Standing Elk for help in translating the name accurately. The name refers to, in part, to the large tubular snout that the animal possessed. The cranium and an incomplete set of teeth make up the type specimen, seen here.

The Macdonald Primate

Type material from Macdonald 1963

This week is rapidly turning into a week dedicated to not only the Sioux language and its influence on paleontology, but also J. R. Macdonald. We have done weeks like that in the past, but it is not necessarily what we are doing this week; it just happens that the animals we are describing this week were named by Macdonald. Today we are discussing one of the only fossil primates known from North America; this excludes primates of Central America. Ekgmowechashala, meaning "Little cat man" in Sioux, is a genus of 5 lb lemur-like primates standing approximatel one foot tall. Macdonald's etymology guide in the description shows that the pronounciationis  Igg-uh-moo Wee-chah-shah-lah. Living during the Late Oligocene and Early Miocene, Ekgmowechashala consisted of two species, E. philotau Macdonald, 1963 and E. zancanellai Samuels, et al. 2015. The genus has been contentious since the type species was described by Macdonald in 1963, but a recent phylogenetic study by Ni, et al. (2016) solidified the phlyogenetic position of Ekgmowechashala as a member of the group Adapiformes, the extinct relatives of lemurs and other basal primates. Adapiformes are some of the most basal members of the primate family tree. The teeth of Ekgmowechashala represent the majority of the originally known material and allowed for inferences of diet. The diet of Ekgmowechashala is thought to have consisted of soft fruits from the warm forests of the Oligocene-Miocene.

Warm forests are not the typical scene for the Rocky Mountains, but during the time of Ekgmowechashala the mountains of western North America were warmer than they are now. The teeth leading to this inference make up a large majority of the fossil material that has been recovered. Teeth have been recovered from the 1960's to the present day from Shannon Count, South Dakota (type material), Pine Ridge and Oglala Sioux reservations in South Dakota, Oregon, Nebraska, and Texas. Ekgmowechashala zancanellai material comes from Oregon exclusively at the moment whereas the type species consists of material from the remaining localities.

Macdonald's Dog

As we mentioned on Saturday, J. R. Macdonald named a large number of mammals from the Oligocene using the Sioux language prominently in his papers. One of these mammals is the hesperocyonid canine Sunkahetanka geringensis Barbour and Schultz, 1935. The name means "Large-toothed Dog" though the Sioux words are actually constructed in reverse order here. The specific epithet refers to the formation from which the specimen originated; the Gering Fromation of Nebraska. Macdonald also provided a pronunciation guide for Sunkahetanka: Sung-kah Hee Tahn-kah. The teeth of this hesperocyonid are actually enormous compared to many other contemporary canids and others within the family tree. The skull is short and wide with robust zygomatic arches and what Macdonald describes as a massive mandible. The massive teeth, jaws, and well built skull lead Macdonald and others to hypothesize the diet of Sunkahetanka to have consisted of anything it was capable of catching or finding. This includes scavenging bones similarly to the manner in which hyenas do now. This is facilitated, of course, by the enormous teeth and the massively built skull in which they are operated. Unlike last week's Leptocyon, Sunkahetanka looked much more like a wolf or a large dog as we now think of them. At a weight estimated (by Legendre and Roth 1988) using tooth size of between 12.8 kg (28 lb) and 13.8 kg (30 lb), Sunkahetanka was about the size of a large beagle (I prefer thinking of basenjis; points to people that know them). This is considerably smaller than a wolf, but imagine a beagle sized animal that could crack and break bones.

©Ghedoghedo via Wikicommons

Oldest Sioux Animals

The oldest fossil animals with Sioux inspired names are technically the variously assigned Brontosaurus or Apatosaurus yahnahpin. A yahnahpin is a breastplate and graced the sauropod's name in reference to the well preserved gastralia, or belly ribs, that were recovered in the fossil remains. These remains date from the Jurassic of Wyoming and the information known about the animal and shared online is mostly in the domain of message boards and academic squabbles. The best description of the material comes from the Wyoming Geological Association Field Conference of 1994, which is, unfortunately, not available online.

Another dinosaur that prominently features the Sioux language is Dakotadon lakotaensis. The name refers to a Sioux tribe, the Dakota, and the Lakota Formation of South Dakota. The name of the formation, of course, refers to another Sioux tribe, the Lakota. Originally named Iguanodon lakotaensis in 1989 by Weishampel and Bjork, the species was reassigned to a new genus as Dakotadon by Paul in 2008. Still an iguanodontid, Dakotadon is a basal member of its family and has been suggested, but not yet shown to be, a synonym for another basal iguanodontid, I. bernissartensis. The arguments continue over the phylogenetic position of the dinosaur, but the basal position is not very contentious. The fossil is important to the fauna of North America as well as being important as a representation of the Sioux language because it represents the only universally accepted iguanodontid from the continent.

Named Through Sioux

The languages of the Sioux Nation have been sampled and featured extensively in North American paleontology. Examples span from dinosaurs such as Apatosaurus yahnahpin Filla, et al. 1994 (currently assigned to Brontosaurus), a rare North American primate, Ekgmowechashala philotau Macdonald 1963, and the anthracothere Kukusepasutanka schultzi Macdonald 1956. Mammals constitute a large portion of the Sioux names in paleontology, but most of those are actually due to the works of James Reid Macdonald (read his 2004 obituary here) who prolifically described and named mammals from the Miocene and Oligocene from Oregon to the Dakotas using the Sioux language. The immense number of specimens bearing Sioux names means that we can easily pick any single animal to celebrate the contributions of the language to paleontology, but I think we can all agree that this would not show that contribution the proper justice it deserves. Instead, for the first time ever, we will discuss a different animal every day this week.

Holiday Over

Back from the holiday (in the United States), we are going to end this canid filled week by trying to find a good image of Leptocyon. There are not many illustrations of Leptocyon. There are no well established reasons that there should be fewer Leptocyon illustrations than there are online, in books and magazines, or in museums. Most of the illustrations that are online that are labelled Leptocyon look like animals that are somewhere between cats and dogs in appearance. Many other interpretations make Leptocyon look like coyote or a fox. The image below, however, shows Roger Witter's interpretation of Leptocyon mollis, a species named in 1906 by Merriam but not officially listed in the genus, from the John Day Assemblage of Oregon. This interpretation looks more like a wolverine than it does a dog or a cat. It is an interesting interpretation of a very dog-like skull.

©Roger Witter

Papers on Dogs

Leptocyon was not much larger than a dachshund, with an estimated weight of approximately 3.26 kg (7.2 lb). The description of many websites stops after mentioning that fact and that its slender jaws and gracile body gave Leptocyon the appearance of a small fox. These are not the only characteristic attributable to Leptocyon however. The initial description and naming of the material is attributed William Diller Matthew in 1918 (see page 7 of the PDF). Matthew noticed the slender mandibles immediately and described the teeth and their morphology in detail. Matthew also states that the morphology of Leptocyon appears to make its inclusion as the ancestor to not only canids but also foxes, unlike some previous authors must have declared. As we have seen throughout the week up to now, most of the sources of information about Leptocyon are sources that describe the entire evolution of the canid family, not just descriptive works of the morphology or paleobiology of Leptocyon. Wang, et al. 2004 is a fairly well-known and (I have not checked into this recently) definitive description of the phylogenetic evolutionary ecology of all canids. It is a book chapter, but for the reader curious about Leptocyon or the dog enthusiast curious about where their pets came from, this is a good read with a lot of detail and a fair amount of technical elements to it.

All Your Fox-like Facts

Leptocyon looked something like a fox, regardless of the species. There are no videos that show this fox-like species as scientific illustrations. There are a few videos that briefly mention Leptocyon, but the only one that is really worth watching is this video on the evolutionary history of canids; the mention is brief but the entire history of canids is worth watching on its own. ThoughtCo (previously About.com) is really the only dedicated website with detailed information on Leptocyon and the site hosts two webpages concerning the animal: one dedicated to the evolution of canids and a second with quick facts and images (and interpretations) of canids throughout history (Leptocyon is #12).

Furry Friends and Megasymbol

Appearing in the Oligocene and peppering the fossil record into the Miocene (from approximately 25.8 - 10.3 million years ago), Leptocyon is a genus of small bodied canids endemic to North America. These canids, consisting of three species, L. gregorii, L. vafer, and L. vulpinus, represent the origins of the canine family tree as it diverged away from the feliniform carnivorans and the other caniforms. The caniforms include not only dogs, foxes, and wolves but also bears, seals and sea lions, and mustelids. For the approximately 15 million years of the fossil record that Leptocyon appears, it is the only canid that appears. Around the time that Leptocyon disappears from the fossil record an explosive diversification of foxes, wolves, coyotes, and wild dog begin to appear in the fossil record.

In the vein of an animal of importance to the native peoples of this continent, this precursor to so many different types of canines is the grandfather genus to many very impactful animals on the mythos and lore of those peoples. Wolves and coyotes are extremely important to many cultures of North America. Foxes, though, are just as important but not as prominently featured in Native American cultures as the other cultures. All of these animals star in other cultures globally (I wrote an honors thesis in undergrad on foxes in Japanese Shinto, for example) as well. Our discussion this week will elaborate on the origins of all of these animals from their humble beginnings in North America. The story of Leptocyon is the story of some of our favorite pets as much as it is the story of animals important to the native peoples of North America.

Finding a Unique Illustration

©Christopher DiPiazza

Almost every illustration that comes up in a search for Argentavis depicts a bird landing or taking off with the few exceptions that show the animal simply stretching its wings one way or another; the bird on a dead animal trying to scare off scavengers is a very common theme. The large wingspan and body of the bird are central to the identity of the fossil, so these themes make sense. One of the most charismatic images that I have found does incorporate the wide wingspan of Argentavis, but it also has a piece of its last meal in its beak. Although almost all of the illustrations of Argentavis already looked fairly fierce, this interpretation, possibly because it was drawn head on, looks more intimidating and angry. As we typically see in Christopher DiPiazza's work, the tones and colors are soft and very pastel-like. Despite this, the details are sharp and the pose is dynamic. Additionally, read the linked blog post by the artist. He has hit a lot of the same points we have hit this week, but his insights into his art shine through in his writing, and they are worth reading about while admiring the work.

Massive Eggs and Wings

The egg of Argentavis is estimated to weigh approximately 1 kg and it has been hypothesized that they were laid once every two years. At 1 kg the egg is only a little smaller than that of the Common Ostrich, but Argentavis' egg laying cycles were similar to gulls and albatrosses rather than animals that reproduce annually. It has been hypothesized that the incubation cycle of these eggs was such that the birds were forced to incubate over winter. Chicks were thought to have lived with the parents for approximately 16 months before permanently leaving the nest. By contrast, the Wandering Albatross is the longest fledging extant bird, with the young bird remaining in the nest for 278 days. As with many extant gulls and other seabirds, Argentavis is thought to have then had a sexually dormant period, not achieving maturity until approximately 12 years old; Royal and Wandering Albatrosses reach maturity between 6 and 10 years. The fact that Argentavis was so large means that most predation and death probably occurred either in the nest or by from accidents and old age. How old Argentavis lived to be naturally is up for debate, but we know that a lot of extant birds, large and small, live extremely long lives today. The Kakapo of New Zealand is thought to live well over 100 years; with so few in the wild and their histories not being cataloged until recently, however, the oldest known member was approximately 80 at his death. Other parrots have been known to live into their 80's in captivity and individual Royal Albatrosses have been documented at 58 years old in the wild. The largest flying birds, Great Bustards, live to approximately 10 years, whereas the oldest eagles have been recorded at between 30 and 40 years old (depending on the species). All of these numbers make pinpointing the ages of large birds, especially those that can fly, difficult. Argentavis could have lived a lifespan like that of a large eagle, meaning that it could have lived up to 40 years. That means that an adult pair, laying one egg every two years, could have possibly reared 14 young during their lifespan. Not only a large bird, Argentavis may have had a rather sizeable population at one time or another because of their long lives, large size, and dedication to a single offspring.

Flying A Great Bird

One of the questions that appears time and again with giant flying animals is "How do they get off the ground and how do they stay off the ground?" Because that is a popular theme with large flying animals, the first hit in a paper search for Argentavis is Chaterjee, et al. 2007: The aerodynamics of Argentavis, the world's largest flying bird from the Miocene of Argentina. The author's conclusions are centered around the hypothesized aspect ratio of the wing and estimated body weight. These parameters lead them to conclude that Argentavis was most likely similar to extant vultures and large condors in that it was probably not capable of sustained powered flight, instead choosing to use thermal soaring as its preferred method of staying airborne. Intermittent powered flapping would have been used as it is by these extant analogues as a secondary anti-stall measure but not as a power source for extended flight. This paper builds off of the new data and goes into more computer simulation than Vizcaino and and Farina1999, which initially tackled the problems of Argentavis flight without computer simulations, instead, it appears, relying on estimates of body size and inferred wing shape and comparing these with extant animals and known aerodynamic principals; the full text is not available anywhere online and what I have inferred comes from the abstract found here. The final article I will mention today addresses ecology (and reproduction). Palmqvist and Vizcaino 2003 details ranges, needed amounts of food, airspeeds, and clutch size to determine the ecological impacts and roles of Argentavis. Instead of spoiling this paper by writing in those facts, as I did above to a slight degree, I am going to simply encourage everyone to read and discover the paper's findings for themselves here. I find the paper to be interesting and find myself wondering if anyone would refute any of these findings; I have yet to find a paper that does so (I admit my search is short right now though).

Quick Facts

Above are some quick facts about Argentavis presented in video form. This very quick video does not reveal much that we will not get into ourselves. There are some cryptozoology documentaries on the internet also; whereas these are strangely interesting, they are not necessarily historically accurate or important to watch in order to gain more knowledge about the bird. Aside from these kinds of videos and the short video shared here, most of the videos that show up with Argentavis as a keyword are related to video games.

Birds and Thunder

©Nobu Tamura

In general, native peoples of both South and North America have a number of legends and mythologies describing giant flying animals. From the Thunder Bird (a generic term that encompasses a wide range of mythological figures from different tribes) to Tah-tah-kle'-ah (a race of cave dwelling "owl women" from Yakama lore) to Achiyalatopa (from Zuni folklore), native peoples have been influenced by the idea of giant birds flying across the sky. In some instances these are giant bird-men/women but regardless of the anthropomorphizing of giant birds, these legends could have been influenced by real living animals. One such giant flying bird has been discussed here before (See Pelagornis). Another giant bird, more often associated with the southern hemisphere but by no means entirely limited to that hemisphere, was Argentavis magnificens (Magnificent Argentinian/Silver Bird). Flying through the skies during the Late Miocene and known, at present at least, from only Argentina, Argentavis possessed a wingspan of approximately 6.07 m (19.9 ft) and 72 kg (159 lb) by the greatest estimates. This makes it the second largest flying bird ever known behind Pelagornis. In comparison to living birds, the largest wingspan is that of the Wandering Albatross at 3.65 m (12 ft) and the heaviest (sustained) flyer is the Kori Bustard at 11.4 kg (25.1 lb). At this size Argentavis must have flown much like Pelagornis; flapping powered flight may have gotten it airborne but thermal soaring would have been the most likely model for sustained flight.

Iconic Images

Probably the best Bison latifrons image I have seen this past week (before looking tonight as I write this sentence) is that of Davide Bonadonna that I shared on the Facebook version of this page. Rather than paste it in here without contacting Davide, I will link his gallery where he hosts his illustrations. A large majority of bison illustrations are not as exciting as Bonadonna's or include modern bison being attacked by wolves (if not a hunting scene with Native Americans. In the "non-action but still fun" category one of the best images I have seen this week is Christopher DiPiazza's B. latifrons at the watering hole. I think I like this image more because it shows an interesting level of curiosity not often associated with the big brutish bison and also because it has softer tones that contrast with a lot of the images we have looked at over the course of the week. Please feel free to share more illustrations that you like in the comments.

Documentaries Everywhere

I was considering what to do about the popular culture references today. There are a number of things that I could do, obviously, as we know that modern bison are something that the public is well aware of. I think, instead, that we should look at some documentaries which discuss the historical significance of bison as well as their current conservation status and situation. Though these documentaries do not specifically discuss the three species we have been discussing this week, the life history of extant bison is similar to that of their extinct family members and is therefore informative. The National Park Service covers a little history and their extant modern herds in the linked page. This second video, from the Kratt brothers (in their Wild Kratts series), may be a cartoon (mostly) but its appeal to the younger audience is not to be overlooked. The information that is given out in this video is useful, accurate, and does indeed go beyond it being a children's cartoon. There are also somewhat more traditional documentaries such as this "Fabulous Animals" show and Facing the Storm: the Story of the American Bison (which is movie length). I know that we are well away from extinct animals with these shows, but being able to see a modern descendant and how it lived on quite similar ground (ignoring the members of B. latifrons and B. antiquus found along the eastern seaboard) is important to trying to reconstruct the lives of the extinct animals.

Buffalo (Bison) Papers

Many descriptive papers have been published for Bison latifrons and B. antiquus. Bison occidentalis, however, does not have as many description papers published concerning its remains. There could be any number of reasons for this, but after looking over papers and fossil locations as well as the Paleobiology Database (PbDb: https://paleobiodb.org/navigator/#/a6d8c0b8) the major reason for this might be that some sources, at least, consider B. occidentalis to be a subjective synonym for B. antiquus. This means that the type of the two species is not a shared fossil and that the debate on their synonymy is open. Unresolved taxonomy like this can lead to some sources like the PbDbnot acknowledging the species as unique while others at the same time do still consider it a unique species, such as Hawley, et al. 2013 which describes the occurrence of B. occidentalis fossils throughout the Wisconsin and Minnesota.

The range of these species has always been an interesting topic for research descriptions and has proved fruitful as well. Representative ranges can be hypothesized for each with articles detailing discovery sites and discussing their significance to the ranges. For instance, B. latifrons has been described in South Dakota (see above), Kansas, Nebraska, and even Georgia. Bison antiquus has been described from remains found in Florida, Alberta, Canada (with a mention of B. occidentalis as well), Washington, and Southern California. Bison occidentalis has been described from sites in Iowa, Alberta, Canada, Texas, and California.

A lot of papers go beyond simple description of these three taxa. A large number of papers published about these papers are from archaeological journals, which is highly logical. Bison, regardless of species, have been some of, and eventually the absolute largest, herbivorous North American mammals encountered by humans. Their use as food, clothing, crafting sources, weapons, and fuel for fires is well documented in North American historical sources and from archaeological digs (as a reference read Agogino and Frankforter 1960 and Wheat 1967). Associations of bison with archaeology is to be expected and some of the articles detail not only interesting aspects of the animal's life histories (refer to this article on butchering and taphonomy as an example) but also a great deal about their capability to endure hardships both human inflicted and natural. These include descriptions of diseases that have been studied in B. antiquus as well the effects of hunting in B. occidentalis.

There are a lot of articles here today, but I cannot apologize for the rich literary record of bison research; nor would I want to if I was asked to honestly.

Three Sets of Information

My plan for this week is to, unlike other weeks, treat the three endemic species of North American bison as individual animals rather than discussing the genus as a single entity and pulling out interesting information regarding individual species. The reason we are approaching the information this way is because, other than overlap zones in the history of the genus, the three species are radically different morphologically and warrant their own attention as parts of the whole evolutionary history of the genus Bison. For a quick overview of fossil bison without reading the following paragraphs a short stop in the digital library of the San Diego Zoo may be all you need. To start, we will look at the three species in order of their history, meaning that B. latifrons is our first stop.

The artist Roman Uchytel is typically mentioned when we look at his illstrations and interpretations of paleofauna. Today, however, his art coincides includes a concise history of B. latifrons. His illustration, of course, is a nice addition to the information, but not the primary reason to follow this link to his website. Not to be outdone by the display shared yesterday from North Dakota, other states also have information on B. latifrons that were unearthed within their borders. Speaking to the range of the animal, there are also finds from San Diego and as far as Florida (B. antiquus is also mentioned on this site, attesting to its range as well) that are mentioned. This site shows and mentions the sale of fossils but such practices are not endorsed or encouraged here.

The intermediate species, B. occidentalis, also has space and information dedicated to it on Roman Uchytel's site. Again, his illustration is a positive bonus to the information provided. Most of the information is taken from Wikipedia, but that is not too important here. Bison occidentalis is not as well known as B. latifrons or B. antiquus, and most of the information online about this species is actually in the form of descriptive papers, which we will save for Tuesday.

As with the previous two species, B. antiquus has been illustrated by Roman Uchytel and he has paraphrased information from Wikipedia. Placing the three side by side is informative and interesting in its own right actually, so it might be worth the momentary effort. Bison antiquus has a few more resources online. This is partially due to its more recent history and of course to those resources being studied a little more in depth than the other fossil species. In addition to Florida and the western part of the North America, there are specimens of B. antiquus known from Ontario, Canada, Texas, and Florida.

November in North America

It is possible that if you are from North America you are aware of the trend in the last 20 or so years to take back holidays celebrating European domination of the native populations of this continent. I remember personally seeing protests at Thanksgiving parades in Plymouth and more and more states appear to be dropping Columbus Day as an official holiday or at least renaming it. Given that the United States of America observes a Thanksgiving event in this month, the same holiday as the protests just mentioned for anyone not following U. S. holidays, I thought it might be interesting to look at the fossil history of some of the groups of animals that have been of special importance to the native cultures that once spread across this continent and that we have, to a significant degree, lost over the past 400 years. This is a look into how the animals that became important to the peoples of North America became the animals that they are now and what their fossil history, regardless of how recent those are, reveals about their evolution. I know these are not dinosaurs, but life on Earth is complex and filled with a lot more than dinosaurs, so hold on for the ride, or take a break and come back to us in December. The first animal that we will look at this month is the America Bison.

The American Bison (Bison bison) has a long history that, like a significant number of animals, starts on another continent; there is an extant European Bison (Bison bonasus) distantly related to and far more rare than the American Bison. The fossils we are going to discuss are the members of the family that arrived or thrived in North America prior to the appearance of what we would recognizably call the American Bison. Three extinct species of the genus Bison were endemic to North America prior to the appearance of Bison bison, the American Bison, in the fossil record. These species were, in order, B. latifrons, B. occidentalis, and B. antiquus. They range from approximately 240,000 years ago when B. latifrons is thought to have initially crossed the Bering Land Bridge into North America to approximately 5,000 years ago when B. antiquus appears to have gone extinct. The three species are not identical and B. latifrons is actually quite impressive, measuring in at 2.5 meters (8.2 ft) and estimated to have tipped the scales at 2,000 kilograms (4,400 lb); that is almost as tall as a modern Asian Elephant. The horns of B. latifrons measured approximately 213 centimeters (84 in) from one tip to the other. Bison bison measures in with horns approximately 66 centimeters (26 in) from tip to tip. Bison occidentalis is less well-known, but it is smaller than B. latifrons and its horns, also much smaller, pointed toward the rear of its skull, rather than the front. Bison antiquus was nearly as tall at 2.27 m (7.5 ft) but much lighter at 1,588 kg (3500 lb) and with horns noticeably smaller, though pointing forward, that measured approximately 1 m (3 ft) wide.

Photo from North Dakota Department of Mineral Resources. B. latifrons exhibit at the North Dakota Heritage Center

All The Pictures

What is the absolute best illustration, scientific image, line drawing, or photograph of a fossil of Helicoprion? If you cannot think of an answer, you are not alone. Is it this image of the estimated size of Helicoprion? It might be this image of all of the hypothesized Helicoprion morphologies (I saved it for my own visual delight).  The Mary Parrish illustration shared on Wednesday is up on the list of good illustrations for most people. There are a number of poor illustrations; we will not share those here today. My personal favorite, as we always see here on Fridays (feel free to share your own of course, but I write it, I choose it typically). The thing this week is that I do not really have a favorite. Some of the best illustrations are outdated and terribly wrong, despite their wonderful appearance. Mary Parrish's interpretation is wonderful, and Nobu Tamura's is as well (see Saturday's image). The image for today, rather than being a favored illustration, is one of my favorite photographs of a Helicoprion fossil. This tooth whorl is immense in appearance and its rolling form is somewhat mesmerizing. I almost cannot stop looking at it honestly.

Other Anatomy

The papers yesterday described the anatomy of Helicoprion's tooth whorl and as such, though we usually pick the most interesting anatomy and describe it on Wednesdays, we will have to discuss something slightly different this Wednesday. The topic for today still has to do with the tooth whorl; it is actually almost impossible not to discuss the only two materials that are known from Helicoprion  fossils (crushed cartilage and dental remains). Two hundred ninety million years ago Helicoprion was a successful predator in oceans globally, but what was it eating with its strange tooth whorl? The tooth whorl, according to the Smithsonian reconstruction, was not the first line of attack for this shark, and that is something important to talk about in terms of diet. When the tooth whorl is placed in the older positions (in the front of the mouth, the middle of the mouth, as an extension of the chin) the scenes we saw in the Animal Planet clip make a lot of sense; Helicoprion slicing fish and squid in half furiously as it swims about in the ocean. The interpretation of Mary Parrish (with help from Robert Purdy, Victor Springer, and Matt Carrano) places the tooth whorl in the gullet of Helicoprion. This positioning places the whorl in the same area as the gills (consisting of five gills rather than six/seven as is common in basal extant sharks for the shark enthusiasts) but without blocking their ability to extract oxygen from the water. The solution to the tooth whorl being in the throat was not having to swallow food before being able to chew it or bite at it. Instead, this reconstruction possesses a set of teeth similar to those of other sharks; a cartilaginous jaw housing rows of teeth that are used for catching, injuring, and otherwise stopping prey from getting away. The teeth in the throat, then, could have been used to snag prey as it entered the oral cavity, with the more traditional teeth pulling the prey into the mouth and the tooth whorl pulling the prey into the throat. Extant animals such as some snakes, fish, and other reptiles as well as extinct animals like mosasaurs possess teeth on the palatal bones that act in a similar way, helping to hold and move prey further into the throat and down the esophagus than mandibular and maxillary teeth are capable of doing. This version of the purpose of the tooth whorl is practical, if not as fantastic as some of the past iterations of the curious anatomical hypotheses.

Papers for Halloween

Is there a better way to spend Halloween than reading papers about a fascinating and scary looking shark? There might be, but I am not going to tell you how to spend your evening. One could spend literally all night reading about Helicoprion; there is seriously that much literature out there on this very interesting shark. I will gladly pick out a few articles for everyone today though. First, I recommend reading both Chorn, et al. 1978 and Eastman 1900 for reviews of the early and more recent reviews of Helicoprion descriptions. Next, I would recommend reading Tapanila, et al. 2013. This study includes a CT image study of the tooth whorl and description of its anatomy and corrected placement in the mouth including a massive illustration of all of the hypothesized placements of the tooth whorl in recorded publications. Lastly, I suggest Ramsay, et al. 2015 which further explores Tapanila, et al.'s hypothesis, CT data, and compares Helicoprion with other genera of shark. This publication also discusses the overall morphology of the shark and how it went about using that tooth whorl in its daily life.

Prehistoric Terrors

Helicoprion has appeared in a number of documentaries and has been mentioned in a number of shows that do not really concern the intriguing shark. This short clip from Animal Planet's River Monsters is the best clip available online from any of the documentary mentions of Helicoprion. It is also the only clip online other than the Discovery Canada clip shared yesterday. As a bonus, one can see a photo of a 3D printed tooth whorl on the River Monsters website for the episode Prehistoric Terror here.

Scaring Children with Teeth

The teeth of Helicoprion can be somewhat frightening in a number of different ways. However, we should probably make sure that the kids in the audience are not frightened when they look at sites like Prehistoric Wildlife that showcase the teeth and the many different hypothesized alignments of those teeth over time. The number of alignments and positions of the teeth are actually somewhat amazing, all things considered; every idea that was put forth had those teeth in some interesting and new position in the body of the shark until their actual position was surmised in the last 50 years or so. In addition to the history of the teeth on Prehistoric Wildlife, there are a number of short informational videos and clips of longer documentaries from Discovery and Animal Planet that are worth checking out. This shorter clip from Discovery Canada goes well with the comprehensive fact page and leave us some other quality videos for tomorrow.

Fearsome Sharks

©Nobu Tamura, 2016; most recent accepted iteration of whorl location

Few animals are as frightening to humans, as primates, as large oceanic predators. The diversity of sharks is such that we could discuss almost any member of the shark family and have a "scary" animal for the final full week of October. What most anyone would imagine is the obvious choice for a scary shark would is something in the Carcharodon genus; that being the group that contains C. megalodon (Megalodon) as well as C. carcharias (Great white shark). Instead, because it is one of the most interesting of the odd members of the shark family tree, this week we will discuss the genus Helicoprion. Contrary to most popular knowledge, Helicoprion is not a single species, but rather a genus consisting of 3 acknowledged and accepted species and 3 questionable species. These species are: H. bessonowi Karpinsky, 1899; H. davisii Teichert, 1940; H. ergasaminon Bendix-Almgreen, 1966; ?H. karpinskii Obruchev, 1953; ?H. mexicanus Mullerried, 1945; and ?H. svalis Siedlecki, 1970. Helicoprion is better known as the "tooth whorl" shark, owing to their unique spiral arrangement of teeth. The shark is known from global deposits spanning North America, Europe, Asia, and Australia, but the cartilaginous elements of the skull, spine, and body are unknown because of their relative inability to fossilize well. The whorl of teeth was, because of a lack of associated material, at one time or another, thought to have been the nasal process, a throat structure, and teeth, of course, placed in different portions of the jaw. Different specimens are named from different regions and for having disparate tooth whorls; the only possible evidence for individual species given the lack of other skeletal elements for most of the specimens.

Iconic Representation

Deviant Paleoart via Creative Commons

As I have mentioned a few times this week, there are many illustrations of Majungasaurus out on the internet for people to view and be entertained by. There are many images of the dinosaur running, such as this one (though it is a bit skeletal). There are also interpretations of Majungasaurus roaring, which look quite frightening, to smaller dinosaurs, and is somewhat stereotypical, but nonetheless, classic subjects of illustration. This version of the mouth open posture is simply showing an interpreted gape and the rows of teeth Majungasaurus had in its mouth, which is nice to see and a nice inset to the main illustration. There is even am illustration on the National Science Foundation page detailing the hypothesized air sacs of Majungasaurus. These are all well done and worthy of note, but whenever I think of Majungasaurus there is one very specific illustration that always comes to mind. The image, shown here, is a specific view of the head of Majungasaurus in high detail. We could spend a day discussing, critiquing, and appreciating all of the illustrations here, but instead, as I am backdating this entry, we will move on shortly.

Majungasaurus Everywhere

Majungasaurus has been seen in many different forms of media for the past 60 years. Originally described by the French paleontologist Rene Lavocat in 1955, Majungasaurus was named as the dinosaur of the province it came from, Mahajanga in northwestern Madagascar. Lavocat's described remains were not the first remains described, but the 1896 description of theropod remains from Mahajanga were published as new remains of Megalosaurus by another French paleontologist, Charles Deperet. Given this extremely long history of remains, the amount of popular knowledge and subsequent popular culture references of Majungasaurusought to be much more extensive than it currently is, despite the mistaken name. Following the 1979 description of remains under the name Majungatholus atopus (Sues and Taquet) the dinosaur garnered more attention, as a pachycephalosaur, until its theropod and cannibalistic nature became more well known and widespread after the 1998 discovery of Majungatholus-like set of remains. Those remains were described and reassigned to Majungasaurus, with the name Majungatholus then synonymized with the older name.

Aside from the two documentaries shared earlier this week Majungasaurus has not appeared in many places on television or in movies. However, Majungasaurus has been illustrated many different ways by many different artists and has been written about many different times by many different authors. Majungasaurus appears in dinosaur encyclopedias aimed at kids, kids picture books, general dinosaur knowledge books, and even college level texts. Perhaps speaking even more to the popularity of Majungasaurus is the copious number of toys and models produced in different poses and from different molds.

Papers Everywhere

Being an ugly cannibal dinosaur has its perks. In the scientific community it has perks at least. The number of papers written about Majungasaurus (including its synonymous genus Majungatholus) is very respectable. The paper regarding cannibalism was shared the other day, though if you cannot find it through an institution of your own it is difficult to find online. I would share it if I had permission to do so, unfortunately it is not my paper. I can point out some other good topics that have been covered though.

The interesting anatomy of Majungasaurus is discussed in detail in a 184 page special memoir entirely dedicated to Majungasaurus crenatissimus published through the Journal of Vertebrate Paleontology in 2007. A number of different topics including the appendicular skeleton, phylogney, dentition, and observed pathologies are discussed in the memoir. Particularly interesting to myself is an article on the craniofacial anatomy. The craniofacial anatomy of Majungasaurus is a defining feature of the dinosaur and is covered in depth in the memoir as well. Majungasaurus is a dinosaur with a lot of bumps and nodules on its skull that cause it to look somewhat terrifying and rather interesting at the same time. These morphological oddities have been discussed in passing a number of times by a number of different authors but Sampson and Witmer tackled the topic head on in the memoir The paper discusses the anatomy of the skull in light of a then newly discovered cache of skull material. This entire publication can be found at this link on BioOne, though access to JVP articles is regulated and available to members only without paying for individual articles. I understand that this does not help anyone access the articles mentioned here if they are not members, but this is a one stop source for Majungasaurus information. Those without access can find a similarly populated list here, but please be aware that the list is 5 years old and some links may not work.

Cannibalism Imagined

Due to the highly publicized knowledge of cannibalism in Majungasaurus there have been multiple iterations of animated and illustrated versions of Majungasaurus engaging in cannibalism. As stated earlier in the week, the act of cannibalism is intriguing to many different people and that is why there are so many different interpretations. There are two different documentaries that have animated this behavior. The two documentaries independently portray active predatory situations wherein one Majungasaurus attacks and eats a second individual. This, as we stated before, is hypothesized and, while making for good television, is certainly not a guaranteed certainty. In the BBC documentary Planet Dinosaur the predatory cannibalism occurs when there is a shortage of food and two young animals are still hungry, prompting their mother to secure some kind of food for the young. The second, History's Jurassic Fight Club, approaches the subject in a similar manner. Instead, a solitary Majungasaurus eating a smaller dinosaur it has killed is confronted by another Majungasaurus looking to steal a meal. Fighting ensues and one of the animals ends up with two meals. The downfall of both documentaries is not in presenting hypothetical situations or imagining dinosaur fights (most dinosaur documentaries love doing these scenes) but instead in the fact that Majungasaurus was an abelisaurid and possessed very unique forelimbs and hands. The BBC version of the dinosaur is closer to the reconstructed skeleton, but is still a little wrong because the arms are still facing forward and in a "want to hug you" sort of posture.

Cannibalism on Display

How do we know an animal that is now a fossil was a cannibal? As with much evidence of cannibalism, even in extant animals, we must look at the teeth, the wounds, and how these two pieces of evidence are correlated. In the past, ichthyosaurs (specifically Shonisaurus and the ichnospecies Ichthyosaurolites) and Coelophysis were suggested to have been extremely cannibalistic in their behaviors. A little over 11 years ago, the cannibalistic tendencies of Coelophysis were analyzed and refuted; the supposedly cannibalized animasl turned out to be a basal crocodile and outside of the body of the second Coelophysis specimen accused. Ichthyosaurolites, on the other hand, is an ichnospecies based entirely on ichthyosaur coprolites that contain the skeletal remains of other ichthyosaurs; the name literally means "ichthyosaur coprolite (fossilized feces)".

From Rogers, et al. 2003. Caudal tail chevron of Majungasaurus. White arrows indicate drag marks and black arrows indicate impressions initial biting marks from Majungasaurus teeth.

In the skull of Majungasaurus, the teeth are characteristically shaped, spaced, and serrated. All of these identifiers have helped paleontologists to recognize the marks of Majungasaurus feeding on a number of other animals from Madagascar. Chief among the animals possessing numerous bite marks from Majungasaurus teeth are the sauropod Rapetosaurus and Majungasaurus itself. No other theropods are known from Madagascar during the 70 - 66 MYA time frame during which Majungasaurus is known to have lived. This was used as the first inference concerning the tooth marks on Majungasaurus bones. This in turn led Rogers, Krause, and Curry Rogers to look at the teeth of Majungasaurus and at the marks that they had left on Rapetosaurus bones in 2003; Curry Rogers had led the description of Rapetosaurus in 2001. The initial bite marks that we see (look at the black arrows) are spaced and shaped identically to the teeth known from the multiple Majungasaurus skulls that have been recovered. The drag marks (white arrows) indicae areas where the serrated denticles have been dragged across the bones.

The first argument that is made here, often, is that the results of interspecies combat might look something like this. However, there are two important aspects of these wounds that make the case for cannibalism more compelling. The first is that these wounds show no sign of healing; the bone would have likely attempted to heal itself, at least, a little, after a traumatic bite in which humerous teeth insulted and scratched the bone deeply enough to clearly score the skeletal material. The second is that these chevrons, and many other limb and vertebral elements that also contain bite marks like these, are inaccessible during combat as they are in areas that could only be (easily) bitten when other elements of the body were exposed either through decomposition or predation.

The remaining question, however, is whether this cannibalism occurred as a result of scavenging behaviors or if Majungasaurus actively hunted members of its own species. The two activities have been documented in various extant species including lions and chimpanzees, so neither would be exceptionally abnormal or unique to Majungasaurus. For more reading on the exact findings of the Rogers, et al. team, read the following paper:

Rogers, R.R.; Krause, D.W.; Rogers, K.C. (2003) “Cannibalism in the Madagascan dinosaur Majungatholus atopus.” Nature, Vol. 422, pp. 515-518

An Old Ugly Dinosaur

©Nobu Tamura

One thing that creeps into horror movies and the Halloween season every year is the act of cannibalism. There are cannibalistic animals throughout the animal world but it is a taboo in most human societies and that makes many of us cringe when we hear about cannibalism in animal groups. Dinosaur cannibalism is rarely documented, but one theropod is particularly well known for its cannibalistic behaviors. Majungasaurus crenatissimus was an abelisaurid theropod and the apex, and possibly only large, predator of Madagascar during Late Cretaceous. At the time Madagascar was already an island separated from both the Indian subcontinent and African continent. As the largest predator on the island Majungasaurus had only other members of its species to truly challenge its supremacy as a predator on the island. Whether these clashes led to the evidence of cannibalism or it was a result of scavenging we do not know. However, Majungasaurus' cannibalistic behaviors and its abelisaurid body plan and often craggy frightening skull morphology make this theropod one of the ugly and frightening fossil animals that deservedly we are discussing during October and during the week leading into the Halloween week.

Portrait of an Ugly Therapsid

One of the best things about very odd animals is that they tend to inspire a lot of interpretations and illustrations because they tend to spark the imagination. Estemmenosuchus certainly inspires fantastical illustrations; Dinocephalian fossils have a tendency to inspire fantastical illustrations because a number of them possess very intriguing and unique skulls. The reconstructed skeleton of Estemmenosuchus is equally intriguing; we will not look at illustrations only today however.

©Dmitry Bogdanov

It is important to note that the realistic nature of illustrations can be affected by the type of illustration we are looking at. Dmitry Bogdanov's style, like Nobu Tamura's, is very soft and often portrays the animal in sterile conditions on white backgrounds; this is not true for all of either artist's illustrations. However, this is not detrimental to the art and, in fact, the implied simplicity of the illustration of this Estemmenosuchus uralensis alows us to more thoroughly take in the entire animal and appreciate the posture, the size of the head, and the stout character of the overall animal. Estemmenosuchus, as we knew before seeing the animal as portrayed here, was a sprawling and squat animal with large canine teeth, which are very visible here. This illustration is labeled as a male animal. The largest canines are used as evidence to support hypotheses of sexual dimorphism in Estemmenosuchus in at least one paper.

©Vladimir Nikolov

More realistic appearances of Estemmenosuchus are as reliant on a stark and bold illustration style as the first is on a softer and cooler colored style. These are most realized in the line drawings that accompany the description papers, but can also be found in the styles of artists like Raul Martin, Dinoraul, and Walter Myers. The illustration included here as a representative of the more realistic appearing (because of its hard lines and high contrast as well as lack of soft tones) was drawn by Vladimir Nikolov. The description of this piece by the artist states that the scene depicts two male members of the genus are engaged in territorial combat. The fierce looking faces and skulls of the animals were apparently not enough to warn one another off from actual physical fighting, as we see in many extant species today.

Sprawling Horned Faces

From Chudinov 1965

Estemmenosuchus has a crown of horns. The crown of horns has been hypothesized to have been used for intraspecific signalling and display short of combat; combat with the horns was probably used as an absolute last resort by these animals. The reason that it would have been used as a last resort is that the horns were massive bone structures. Unlike antlers, horns are composed of bone and insult or injury to these structures can be much more traumatic to the animals than damage to antlers (injuries to antlers are serious of course though). The horns of Estemmenosuchus were composed of extremely thick outgrowths of the frontals and cause the skull to appear even more massive than it is. Known skulls of Estemmenosuchus are approximately 65 cm (26 in) in length. That is not the only thing that is large and unique about Estemmenosuchus though. This large therapsid (approximately 3 m  or 10 ft long) also had a sprawling posture; this is somewhat typical in therapsids and Permian reptiles as well. Some have used this sprawling posture as evidence for an herbivorous diet, saying that the sprawling posture enabled the animal to hold a large fermenting gut with more support than if it had a posture like cattle or a similar mammal; this seems less than ideal given what we know about extant mammals. The canines of Estemmenosuchus are used as evidence to a different, more omnivorous but not quite carnivorous, dietary regimen.