Fossils and Extant Crinoids

Videos of extant Crinoid groups are often given fantastic titles like "Amazing Free Swimming Feathers!" and descriptions like "mesmerizing video". These videos are not heavily inundated with facts and interesting trivial bits as we try to disseminate on Sundays, but there are plenty of dedicated websites to garner facts from. Fossil Facts, Fossil Era, and Kids Search are prime websites for reading short paragraphs loaded with all kinds of facts about these interesting animals. More official sites also present fact files and quite a few photographs of fossil specimens. These fossils come directly from Kansas, in the case of the Kansas Geological Survey and a variety of localities in teh case of the UCMP at Berkeley.

Overlooked Animals

One of the most populous and overlooked animals in the ocean since at least the Ordovician is the group of over 500 extant species in the genus Articulata. Also known as Crinoids, these strange animals belong to the class Crinoidea, live in areas from the shallows to at least 9,000 m deep, and have a historical range in three additional genera (all extinct) that spans the globe leaving fossils in a large range of geographic areas and geological time zones. The fossils of these passive filter feeding animals have been found attached to fossilized driftwood, the bottoms of ancient oceans, and in lengths in excess of 40 m. As single stalked echinoderms that were enormously successful, Crinoids underwent two explosive radiations early in their evolution in both the Ordovician, when they first definitively appear in the fossil record, and during the very early Triassic, after an extinction event near the end of the Permian initially bottle-necked the Crinoid record for a short period. This second radiation lacked morphological diversity previously seen in Crinoids, but did exhibit some of the longest individuals with Pentacrinites reaching lengths of approximately 40 m (130 ft).

Given that there are hundreds of species and somewhat fewer genera, the reason that Crinoids have been lumped together for this week is because the wealth of information for any one individual species is actually quite limited, online, and there are a number that are not even mentioned anywhere online aside from acknowledgement that they existed. Talking about the entire group together we can discuss a large number of morphologies all at once and compare them. Be prepared for a lot of talk about stalks and plant-like morphologies. These are very unique animals that are often mistaken for plants. Be amazed, or perhaps underwhelmed, by their looks!

Photo uploaded by Berengi; GNU Free Documentation License, Version 1.2

Today is Something Different

The diet of cave lions was much like the diet of modern lions in that these were large apex predators capable of subduing and preying upon equally large animals. Some of these lions have been found in the dens of bears, suggesting that they were capable of taking bears but that some may have perished in the act; conversely, these remains found in bear caves may have been successful and commandeered the cave for itself. Other known prey items included bison, reindeer (as seen previously), and even saiga antelope; a family of Eurasian antelope that possess rather well insulated nasal passages. Despite never being mentioned, the human imagination likes to imagine that all apex predators would like to get a good meal of human flesh and therefore there are indeed illustrations, like the one below, that shows a "caveman" returning with dinner to find a cave lion devouring his female partner. Be very careful in showing this image as it has no real basis in the evidence we have of cave lion diet or their interactions with humans.

The Mythology of Lions

Lions, cave or otherwise, have been revered as royal, strong, proud cats for a very long time in the human world and extending some of the myths, legends, and beliefs that are lion-centric to cave lions is not a far stretch of the imagination. One of the early standouts of human literature is the Nemean Lion which was said to prowl Nemea in Greece and is within the historical range of both cave lion and modern lion but only the temporal range of the modern lion. Cave lion remains may have influenced the story of a lion so large and fierce that only the great Heracles could stop it from terrorizing the Greek peninsula. Origin stories in the myths of ancient Greece credit the abnormally sized lion as having been the offspring of titans or having originated and then fallen from the moon. Accounting for abnormal size in such ways certainly suggests that the story's antagonist may have been based on sub-fossil evidence of a large lion. The additional story detail that the animal lived in a cave could suggest either creative story-telling or literally finding the remains of a cave dwelling lion (a common occurrence from which the common name of Pathera leo spelaea originates).

Prior to the Greek, Persian, and classical Chinese (to mention a few cultures) mythos of lions, Eurasian Cave Lions were depicted in early cave art. The American Cave Lion has been depicted on various media in native North American cultures in various forms. Many of these appear more cougar-like (the North American cougar or puma has been referred to as the American Lion by many different source) and may actually be confused with the other cat. Considering that these more recent works were likely created in the likeness of cougars, the depictions of the American Cave Lion may actually be lower in number than the internet would have anyone believe.

Panel of the Lions, Chauvet Cave

Geography of Lions

Extant lions are restricted to the African continent in the modern day. The extant populations of lions representing a number of recognized subspecies are dotted across the continent in areas that represent a very minor proportion of their historic range across Africa, Southern Europe, and Southwestern Asia. If we were to consider the cave lions as subspecies of the extant lions, as many have described their phylogeny, then the historic range of lions actually covered the majority of Europe, Asia, and portions of North America. We know the extent of their range because Eurasian Cave Lions have been recovered in various places and are well represented in cave art in a variety of locations across Europe and Asia. The North American subspecies (hereafter today I will refer to them as the American Lion) has been recovered in similar ways: as fossils, from glacial deposits, and a large population has been recovered from tar pits as well. The American Lion is known from a shorter temporal range than the Eurasian Cave Lion (340,000 - 11,000 years vs. 370,000 - 10,000 years respectively) and is considered endemic to North America. The American Lion, despite a smaller range and a shorter temporal span, was approximately 25% larger than extant Lions and 15% larger than Eurasian Cave Lions (which are in turn 10% larger than extant Lions) making it the largest of the lions, including the closest common ancestor between American and Eurasian lions, Panthera leo fossilis. This also means that the American Lion is, as many of these lions are, one of the largest known cats in the history of known felids. The size of the cat enabled it to take prey like the reindeer shown below, usually taken in the modern age by large bears or packs of wolves, for example.

©Heinrich Harder

Genetics and Lions

Ice Age animals have a long, in the somewhat short time that it has been available, history of genetic and isotope research. The reason for this is that the thawed cubs mentioned yesterday have generated not only a lot of sensation but have provided scientists with intact remains that preserved the majority of the soft tissues and organs of the animals. This of course has led to chemical, microbiological, and genetic work being carried out using the preserved tissues of the animals. Some of these studies include molecular phylogeny (Burger, et al. 2004), diet inquiries using isotopes (Bocherens, et al. 2011), and studies of genetic diversity (Ersmark, et al. 2015). Some might be asking "How does one study diversity from a single litter of cubs?" and this is not without its caveats and misgivings; however, it is important to note that we have not discussed all of the populations of known cave lion with some tissue preservation. Many American Cave Lions have been recovered from the tar pits of Rancho La Brea in Southern California and these could add to the genetic diversity of known populations. However, the destructive nature of tar pits has hampered DNA recovery but that is not to say that there have been no efforts to recover DNA. These efforts have been largely unsuccessful unfortunately. This leaves us hoping there are other soft tissue deposits to back up these studies on genetic diversity that have not been mentioned here this week. There are, in fact, soft tissue preserved specimens from across Eurasia and many known from Alaska and Northwest Canada that are mentioned in the paper linked above. These are not entire cubs as we saw yesterday, and are therefore less famous, but are nonetheless quite important in terms of genetic studies of cave lion populations and diversity.

Non-genetic, chemical, or microbiological study is also important in the history of cave lions. These have included phylogenetic studies based on skeletal tissue, specifically cranial and dental characters (Sotnikova and Nikolskiy, 2006), as well as a detailed timeline of the extinction processes that affected and eliminated the Eurasian and American (Stuart and Lister, 2011) Cave Lions from their respective habitats. Though we have been discussing the American Cave Lion in conjunction with its Eurasian relatives seamlessly, there is mitochondrial evidence (see the linked molecular phylogeny paper above by Burger, et. al.) that this population split early enough in their evolution that it has been argued that these represent merely a "lion-like cat" with a very popular misnomer. This phrase was first prominently used in 1969 by C. R. Harington as far as I can tell, but his paper suggests that the North American and Eurasian animals were likely conspecific (Harington, 1969).

Monday's Movies

One of the most important discoveries in the past few years was the retrieval of two frozen Eurasian Lions after glacial melt in Siberia. This discovery has led to discussions about cloning and the paleobiology of the animals. Cave Lions have also been recreated and animated in documentaries. These include a lot of different documentaries actually, but the one linked here (from Walking with Beasts) is shown stalking a mammoth, and is fairly compelling to watch.

Things to Know About Cave Lions

A lot of people love Cave Lions as well as extant lions. This has spurred a lot of websites to host information on Cave Lions and even a 20 minute video which was self-produced that discusses what one amateur fossil enthusiast knows about the large cats. This video is spoken over a few artistic recreations of a few populations of Cave Lions. There is also a shortened video from WizScience that goes over a shorter version of some of the same information. Fact pages that we visit often are also well-represented with Prehistoric Wildlife and ThoughtCo (formerly hosted on About).

Subspecies of Lions

The list of subspecies attributed to lions actually encompasses some of the fossil species of lion that we are going to discuss this week. Rather than discuss the three distinct groups of cave lion separately (Panthera leo spelaea, P. leo fossilis, and P. leo atrox), we will lump the three species together in the sense that we will discuss them as individual populations distributed globally and officially labeled as subspecies of the extant species of lion. "Cave lion", used as a generalized term, refers to European (P. leo fossilis), Eurasian (Panthera leo spelaea), and North American or American Lion (P. leo atrox) populations of large carnivorous felids of the Pleistocene. These animals were not excessively morphologically disparate from extant lions, but distinctions can be seen and observed between the extant and the extinct. These small distinctions add up to larger lions than the extant species, as seen below.

©Roman Uchytel

A Famous Archosaur

Lagosuchus is well known throughout the paleontology and throughout the popular dinosaur and fossil enthusiast circles. The general public may have heard the name of the archosaur, but not as regularly as any other fossil and less than many more. Despite this unsteady popularity Lagosuchus has appeared in a number of different popular streams, books, and cartoons. These include a brief appearance in the original The Land Before Time, though it appears as an announced and non-descript small reptile running around the sauropod next in the opening scenes, but is radically out of temporal context. This version of the reptile is also a victim of the 1980's illustrative process that made most dinosaurs look emaciated and a terrible green color. Lagosuchus has also appeared in a number of texts and popular paintings. Possibly one of the most well-known paintings is one by Douglas Henderson that still looks very lizard-like. This painting is shared on Henderson's website, though it has also leaked onto the internet elsewhere, unfortunately.

Incomplete Remains

The known remains that are still attributed to Lagosuchus are comprised of a single hind leg, a few vertebrae, and a scapula. The hindleg is gracile and slender, indicating that the basal archosaur was lightly built and possibly well adapted for a running life. The vertebrae and scapula are not as indicative in the life history as the hindleg appears to be, but are important in describing the animal. These elements when analyzed together indicate that Lagosuchus was actually capable of running bipedally for short periods of time only, being a facultative biped. Quadrupedality appears to have been the most basic form of locomotion for basal archosaurs, making Lagosuchus a typical early archosaur.

Descriptions and Opinions

Lagosuchus remains, questionable and otherwise, have been described and analyzed many times over. Due to the controversial nature of the remains and phylogeny of Lagosuchus there are not many articles on the animal aside from descriptions of the original materials. These are not hosted online in easily accessible ways because they are slightly older documents from the early 1970's. New material descriptions are available online however, allowing us to look at the known materials and understand how much Lagosuchus material is known to science. There are a lot of papers that use Lagosuchus and the family that was named after it (Lagosuchidae) to describe and discuss archosaur phylogeny. Key in these papers are articles discussing the ankle joint and whether or not there is even any consensus on what the ankle and the accepted basal archosaur tree mean to overall archosaur phylogeny.

Quick Lists of Facts

Lagosuchus, dubious name or not, has a lot of pages dedicated to it online describing the animal or relaying short lists of facts. Today I will simply share a video that summarizes what many of these pages summarize over and over again. It is a somewhat less exciting video than we might normally share for facts, but it does discuss a lot about what we know of Lagosuchus.

Early Archosaurs of the Triassic

Pavel.Riha.CB at the English language Wikipedia

Lagosuchus talampayensis is an early archosaur recovered from the Triassic rocks of Argentina. The Chañares Formation represents the Middle Triassic (230 MA) and, barring recent publications until their results can be independently verified, the some of the earliest remains of archosaurs representing the earliest dinosaurs. Discovered in the 1970's and initially described by Romer in 1971, it has been described by many as a nomen dubium by many. A second species was reassigned in 1994, making the description and known material to be even more questionable. That material is somewhat minuscule in nature as well, which does not help. Instead, we will look at what is known about Lagosuchus, its relatives and what its relationships mean for the current phylogeny of archosaurs.

Everyone's Arkansaurus

There are only a very few widely circulated recreations and illustrations of the little known dinosaur Arkansaurus. The majority of these are variations on a dinosaur that looks very much like the Gallimimus that ran around in the original Jurassic Park movie. A more modern version of the illustrations by Brian Engh does exist that not only updates the general look of Arkansaurus but also adds feathering and a little weight.

Short News Piece Day

Movie Monday is not going to have a movie or a documentary today. Arkansaurus was not, and is not, famous enough to have influenced any documentaries or movies despite its status as a symbol for a state. The only videos about the dinosaur are news coverage of the state's legislative debate over adopting the symbol. There was not a lot of debate actually, but it is a nice little news piece to watch.

Arkansas Geological Survey Facts

The Arkansas Geological Survey (AGS) has a large number of publications and fact sheets describing the state dinosaur and other geological wonders of the state. As a somewhat obscure dinosaur, there are few references online that are more popular. However, aside from sites like Cool Dino Facts and the AGS there are also Arkansas based sites that discuss Arkansaurus. These include Paleoaerie, an evolution and education resource site based in Arkansas (AERIE = Arkansas Educational Resource Initiative for Evolution). Another important site hosting a nice post about Arkansaurus today is Arkansas Life Magazine.

Legislative Dinosaurs

Photo provided to Arkansas Geological Society by ReBecca K. Hunt-Foster

In the past few months there have been a lot of dinosaur stories circulating in the popular news and one of the most relevant to the middle of the United States is the announcement of a new state dinosaur. That dinosaur is named Arkansaurus fridayi and is known from a few bones found by a farmer searching for a cow. In 1972 a man named J. B. Friday was searching his farm in southwest Arkansas and stumbled, almost literally, over a fossil dinosaur foot in the remains of a road cut, which is problematic. Any remaining fossil material may have been destroyed in the construction efforts, by water flow before or after burial, or scavenging prior to burial. The remains were initially described by members of the 1973 SVP meeting as an Ornithomimus which would represent the oldest member  of the species ever recovered. The animal remained informally named Arkansaurus fridayi for 30 years before it was re-described by ReBecca K. Hunt-Foster (published as an abstract under her maiden name; ReBecca K. Hunt). Considering this is the only publication of any type describing these dinosaur remains we will refer to the animal this week with italics rather than quotations; a difference typically separating formally described and accepted genera and species from those used informally. This remains a controversial name, but in defiance (or disregarding the informal nature of the name) the state of Arkansas recently passed a bill making the only dinosaur material known from the state the state's official dinosaur fossil. There is more significance than just naming Arkansaurus the state dinosaur; Arkansaurus is now the 25th official state symbol and Arkansas was the 25th state admitted to the United States.

Too Much Titanoboa

©Jason Bourque

To finish out the past week we need to look at some of the beautiful paintings and sculptures of Titanoboa. The artistic interpretations and scientific recreations of the snake show the animal in typical snake postures and some are stunningly detailed. One of the more tame interpretations of the snake actually shows Titanoboa participating in one of the most common behaviors of snakes or any other animal: casually sunning itself or possibly moving from one place to another. A large dryosaur and turtle help populate the background of this painting and dryosaurs actually figure into more than one Titanoboa representation. The most well known sculpture, the Titanoboa featured in the Smithsonian exhibit for the giant snake, shows a Titanoboa in the process of swallowing a dryosaur. The dryosaur's distinctive tail is sticking out of the snake's mouth as it aligns itself perpendicular to the ground. This seems odd for two reasons: 1) the post hoc knowledge that the hypothesized diet of Titanoboa is composed of fish almost entirely and 2) lifting an animal as large as a dryosaur vertically into the air to swallow it would, it seems, be extremely taxing and difficult even for an enormous snake like this.

Photo by Ryan Quick

The Charmer Snake

Through no mistake has Titanoboa become a popular fossil snake. The sheer size alone is enough to make people ooh and ahh over the giant reptile and the popularity that followed is entirely expected. This popularity can be seen in the number of sites online hosting facts and images of the snake as well as the fact that the snake had a one hour documentary dedicated entirely to the single taxon. In the vein of many internet speculation sites, the Smithsonian's work on Titanoboa even allowed for the fairly ludicrous time traveling fight between a Tyrannosaurus and Titanoboa, giving into the popularity of online forums that pose "what if" encounters between apex predators. We now hypothesize that Titanoboa was piscivorous and not really an apex predator, so many of these sensationalist popular views can be thrown out anyhow. thankfully, without linking a lot of books, we have other popular outlets. The most visible of these is video games and, again, ARK manages to include more fossil animals in the game. ARK tends to err on the side of intrigue ever so slightly when re-creating fossil animals and with Titanoboa the designers managed to find something, even on a snake, to indulge their creativity and the fantasy of the game. The snake looks surly and vicious and manages to do so sporting an almost hairy looking frill:

The Long Remains

Jonathon Bloch with Titanoboa and Anaconda vertebrae. ©Natural History Magazine

The finding of Titanoboa was significant for vertebrate, South American, and herpetological paleontology to name just a few areas and disciplines that benefited greatly from the discovery. Climate studies were another area that has been mentioned a great deal here in the last few days. To truly understand how large the snake is though requires a significant amount of knowledge of extant snakes and how their morphology relates to the size of their overall body. In order to estimate size and weight of extinct snake biologists need to know how the bones and bodies of living snakes are related and then apply the resulting ratios and formulae to the recovered extinct material. Vertebrae, such as those shown from Titanoboa and a 17 foot Anaconda, are often useful in estimating length and weight of fossil animals. Titanoboa was found with a significant number of these vertebrae, which can be rare for snakes. What made estimating the size of Titanoboa easier was not the number and completeness of vertebrae, bones of the spinal column that can make estimating length a bit easier, but the recovery of the skull of the snake. A snake's skull is rarely recovered in extant animal remains, let alone in fossil remains. The reason for this is that snakes are animals with extremely mobile heads. Their heads are capable of moving to extreme degrees and they do so by having very interesting connections between the bones of the skull that are not so much loose as they are workable and even supple. This often leads to the skulls of snakes simply falling apart as the flesh, muscles, ligaments, and tendons decompose and water, wind, and animals scurry about moving the bones. The skull alone is, therefore, a very intriguing find. That makes a very intriguing snake with a very rare skull and an immense body that has captivated millions; not a bad find in a Colombian coal mine.

Titanoboa on Paper

Titanoboa has been the subject of many different sorts of articles from pure description to interesting paleoclimate discussions. The hypotheses concerning life history of the snake are also of particular interest to scientists that want to better understand the enormous boid. The most important papers are those concerning climate. These studies have revealed important aspects of past climates and their changes in addition to revealing clues about how cyclic weather patterns can be used to predict future climate patterns. Weather enthusiasts and snake enthusiasts alike should enjoy these two articles tonite and have plenty to investigate about their favorite topics.

The Monster Snake

Titanoboa generated a lot of interest when it was first described and the illustrations and recreations were first unleashed on the world. In 2012 the Smithsonian Channel released a documentary about the giant snake. The 90 minute documentary still airs, now and again, on the Smithsonian Channel and the schedule is kept up to date on a website for the channel's shows. The Titanoboa page also contains teaser trailers for the show, random clips, and even a game. There are numerous other videos and, were we able to link a free version of the documentary, we could watch that today. Renting it online costs $2, which is not awful, and it is entirely worth the price.

Snake Facts

Everyone loves good facts about fossil snakes. The evolutionary history of snakes is actually coming to light in recent years and recent studies which is of great importance to what we know of both fossil and extant snakes. We actually have a great deal of information about Titanoboa, which is not entirely rare for a given snake, but is less often an occurrence than we typically see in many dinosaurs or fossil mammals. The number of websites hosting facts about Titanoboa are extremely numerous; most are filled with only scientific fact, but there are so many that fact checking all of them would be nearly impossible. Websites we use here a lot, like About (About's dinosaur pages now reroute to a site called ThoughtCo) and Prehistoric Wildlife. New sites that we have used sparingly so far like New Dinosaurs also have facts on the giant snake, despite Titanoboa not being a dinosaur. Dedicated snake websites also share facts about the giant boid like the aptly named Snake Facts site. There are also numerous videos that share information, facts, and animated recreations of the snake that are of great educational value; however, some of these will be saved for Monday.

The Gigantic Snake

The Titanic Boa is not necessarily a clever basis for a name, but Titanoboa cerrejonensis (specifically named for the Cerrejón Formation from which it was recovered) is exactly what its name implies: a titanic boa. The largest known snake (fossil or extant) was recovered from the coal mines of the Colombian region of La Guajira, a peninsula in the extreme northeastern portion of the country. An estimated 12.8 m (42 ft), Titanoboa was thought to weigh approximately 1,135 kg (2,500 lb); the largest extant snake is the Green Anaconda (Eunectes murinus) has an average length of 3.7 m (12 ft) and average weight of approximately 30.8 kg (68 lb). Titanoboa is hypothesized to have been largely piscivorous which would be unique among all boids, fossil and extant. As a boid, Titanoboa would have wrapped itself around its prey and killed using mechanical constriction. As an enormous snake, it would have been coiling itself around its prey to constrict it, meaning that the fish it was constricting must have been equally large, perhaps even titanic (could not resist, sorry).

©Vlad Konstantinov

Quality Tapir-like Fire Beasts

Robert Bruce Horsfall, 1913

There are surprisingly a lot of images of reconstructions and interpretations of Pyrotherium posted online. As one of those fossil mammals that is slightly less popular, it is fairly interesting that the illustrations are as numerous as they are. There are illustrations like the first one that represent the truly old guard of illustration; a time that was powered as much by fantasy as it was by scientific interpretation. We can admit that the scientific interpretation of the day was not quite up to the same standards as modern scientific interpretations are. More current illustrations and interpretations have appeared here numerous times this week and have looked quite dynamic and plausible. Horsfall's illustration is not entirely unrealistic, but compared to the modern illustrations it is a little off from something we could accept as a real live animal. In contrast, and I admit the surroundings of this image help its realistic plausibility significantly, Roman Uchytel's version of Pyrotherium standing in a calm river bank patch of grass looks much more like an animal we might happen upon while walking in a somewhat wild area of the modern world.

Pyrotherium's Head

The head of Pyrotherium is illustrated with a short trunk or a long nose, depending on one's particular definition of the word trunk. That appendage is not preserved on any of the known skulls of course, but the skulls do tell us a lot about the life history of the animal and its cranial ornamentation. The most prominent feature of the skull are the mandibular and maxillary incisors. Two pairs of maxillary incisors and a pair of mandibular incisors are enlarged and elongated as tusks similar in shape to those of a warthog. These give the appearance of the skull tapering and looking fairly sinister, which may have been the intention of such amazing teeth. Additionally, the teeth could have been used to root and dig for preferred food items or to signal health and well being to potential mates.

©Richard Haddad, Beneski Museum of Natural History

Pyrotherium on Paper

The number of articles on Pyrotherium is actually impressive considering the small amount of attention it has garnered online and in the general populace. Description of new species and important revisitations of the skulls have proven immensely important to the knowledge base of Pyrotherium. There is an absolute wealth of information beyond these two papers of course. The entire list of papers is far too long to even bother linking the Google Scholar search list. There are old lectures and a number of "initial" descriptions.

The Statue Videos

Not much of an actual fiery beast, Pyrotherium is lackluster in both video appearances and popular knowledge of the animal. This unknown factor of course inhibits video appearances in most fossil animals and the circle unfortunate circumstances continues from there. However, there is apparently some statuary depicting Pyrotherium in a fossil animal park/garden of statues and this interpretation has been posted online. This statue also roars, apparently, and it is described as "life-like roaring". I will let the viewer be the judge on this. This video of the statue comes from before it was placed in the park:

Fire Beast Cometh

Fun facts abound today that describe and illustrate the life of the stock mammal Pyrotherium romeri. Its name means "Fire beast" and the discovery of its remains in volcanic deposits makes the name relevant, though not in a way that is bizarre or science-fiction like. Somewhat resembling a rhinoceros in stature and a tapir or small elephant in shape, Pyrotherium is a South American ungulate found in Early Oligocene deposits. As with many mammals, the teeth and skull of Pyrotherium are extremely well known and many specimens have been discovered and studied. We know that the incisors of the animal developed into tusks in both the upper and lower jaws. Two pairs of tusks were found in the upper jaw, and one pair were found in the lower jaw. Illustrations and interpretations of the animal bear this out and the teeth are described on sites like Prehistoric Wildlife and Dinopedia (despite the animal being a mammal).

©Nobu Tamura

A Beautiful Parakeet

There are a lot of illustrations and models of Carolina parakeets all over North America in museums and hanging on walls. The internet has even more copies of both the illustrations and the models posted all over the place. Deciding what the best image of Conuropsis is online could take ages and, unlike some of the images we have posted in the past for dinosaurs and other fossil animals, there are none that stand out automatically because they are overly dynamic or show excessive action. One of the few very illustrations that draws the viewer in with action and movement is Michael Rothman's 1984 and 2010 paintings of groups of parakeets flitting about some non-descript trees. One of the more "cute" illustrations Kristina Knowski's watercolor Carolina parakeets nuzzling one another. Possibly my favorite painting of Carolina parakeets is Walton's Ford reimagination of the classic painting the "Death of General Wolfe". If you have not seen the original, please look it over, but  either way, enjoy Ford's interpretation:

©Walton Ford

Eradication or Extermination

Scientific studies of Conuropsis were never conducted or published. However, occupying the forests of the middle third of North America was a medium sized parrot weighing approximately 3.5 ounces, about a foot long with a wingspan approaching two feet. Their green bodies and yellow and red heads could be seen bouncing and flying through the tree tops of deciduous forests of North America, a somewhat abnormal depiction of parrots as we often think of them as rainforest dwelling birds that we see in cartoons and movies. Like most parrots, Carolina parakeets ate seeds, making them a small nuisance for farms. How small a nuisance? Carolina parakeets, at the height of their population, were estimated to have had a population density of up to 2 birds per square kilometer with an estimate of their Florida population numbering in the hundreds of thousands. A farm of 100 square kilometers could have had anywhere between 50 and 200 Carolina parakeets foraging seeds and decimating crops at any given time; the higher number would have been more likely given the supposedly gregarious nature of the birds. Farmers turned to the easiest solution they could find to lower the animal's numbers: destruction of the forest and en masse killing of the seed-stealing culprits. The hunting was not only undertaken to lower the numbers of parakeets foraging in farmland, but also to enhance women's hats. Many birds once suffered this fate, but few have become completely extinct because of fashion.

Books on Parakeets

The definitive guide to the Carolina parakeet, Conuropsis carolinensis, is most likely one of the many books that have been written about the extinct bird. There are scholarly articles as well but the books have done a great job of discussing and describing everything about the bird, its life history, and its habitat. The newest book is a 2004 Snyder book called The Carolina Parakeet: Glimpses of a vanished bird. The book is comprehensive and well worth the cover price. Articles worth reading include the Kirchman, et al. 2012 phylogenetic relationship paper describing the familial relationships of Conuropsis. Because I live in Missouri I find the McKinley papers of the 1960's (The Carolina Parakeet in the upper Missouri and Mississippi river valleys and The Carolina parakeet in pioneer Missouri)to be quite intriguing. There are a number of similar papers for different regions times in history describing the world of the Carolina parakeet.

Single Documentary

As stated yesterday, there is a sad lack of Carolina parakeet material online. There is only a single documentary available online (and I cannot think of any I have seen on television) that discusses Conuropsis, its life history, and its disappearance from the wilds. The 5 minute short looks to be a little bit older, but its quality is acceptable. The scientific quality of the information is also acceptable. Rather than continuing to go on about it, we ought to just watch it, because it was the only thing online to watch this week.

Recently Lost, Quietly Remembered

There are not a lot of sources for the Carolina parakeet. This is perplexing and a little sad given that the Carolina parakeet was determined to be extinct less than a century ago. It is also alarming that an animal as beautiful as the Carolina parakeet could be both the only parrot and a beautiful bird of North America and be largely unknown less than a century later. Extinct animal websites have not forgotten Conuropsis as we can rely on websites like About to bring us facts that are easily readable by most readers online. A slightly higher reading level can be found on the site ExtinctAnimals. The site New Dinosaurs sits comfortably between the two in terms of reading ability and adds in a significant number of illustrations of the bird from a diverse group of artists.

A Subfossil Week

©John James Audubon

On infrequent occasions in the past this blog has focused on animals that are technically labeled as "subfossil" meaning that the animals are recently extinct and are known from remains that have not been buried long enough to undergo fossilization. This week is dedicated to one such animal, a small bird from north America known scientifically as Conuropsis carolinensis Linnaeus 1753. Colloquially known as the Carolina parakeet, this small colorful bird was deemed extinct in the wild as late as 1939 with the last breeding animals dying in the Cincinnati Zoo in 1918; eight years after the last known wild sightings of one of the birds. Once known to range from New England to Colorado and south to the Gulf of Mexico, the Carolina parakeet was the most northern ranging parrot and the only parrot native to North America outside of Central America. As with many parrots, the Carolina parakeet was a victim of the pet industry and its feathers were used for fashionable hats. Additionally, hunting and grazing controls were employed against the flocking bird with no real regulation, allowing for mass hunting like that which caused the extinction of animals like the Passenger pigeon. A great loss for the wilds of North America, the Carolina parakeet is very recently extinct, but little is actually known about the birds, as we will see throughout the week.

Scenes of A River

Some of the best scenes are murals and I thoroughly enjoy the immense extended scenes that murals present to the viewers. Today's artistic interpretation of Stupendemys appears as a character in an elaborately detailed mural-like piece. Clearly this image comes from a book, but would work as a mural (also calling this a two page spread is less flattering than calling it a mural). The Miocene flora and fauna around Stupendemys are all of an older style. The art itself and the coloring also betray the age of this piece; there is no attribution accompanying this image online. Regardless of its age, the scene is well presented if not fanciful, considering the massive amount of animals at the river's edge. The waterway shows a considerable number of the animals we know lived with Stupendemys as well, something not seen outside of mural style illustrations of an animal and its environment.

Popularly Stupendous

Stupendemys is the largest turtle that ever lived, as far as we know as yet. This has not gone entirely unnoticed by fossil animal enthusiasts, paleontologists (professional or amateur), or by the general public. However, the sources for popular culture references are still somewhat on the shallow end of the pool for this amazingly large turtle. The only film reference to the giant turtle is from the straight to video Land Before Time V; I try to not acknowledge that there are movies beyond the original, but they serve as introductory videos to dinosaurs for some very young folks and that I am okay with. Probably one of the best places online to get information from a more amateur paleontologist sort of voice is this 2012 Lord Geekington (Cameron McCormick) blog post. Posting about a blog post in a blog post is a little strange, but given the lack of books and most other pop culture references, it seems a fitting way to end the post.

Turtle Anatomy

As mentioned yesterday, there are two major divisions of turtles aside from binning the animals as freshwater, marine, or tortoise. There are other aspects of the anatomy of Stupendemys that are just as important as the way which it retracted its neck and head. Most of the anatomy that has been described is related to the shell of the turtle as the enormous shell was most readily preserved and fossilized. Additionally, materials such as the humerus, pelvic girdles, and an incomplete set of vertebral elements have been found and described. The "problem" with this turtle is that its anatomy is not that much different from modern turtles as to point out anything exciting or overwhelmingly primitive about Stupendemys. The turtle is remarkably enormous though, with the humeri measuring 310 cm and 340 cm in S. souzai and S. geographicus respectively.

Photo by Ryan Somma at American Museum of Natural History. CC BY-SA 2.0

Types of Turtles

There are two types of turtles in this world, marine and freshwater notwithstanding: pleurodira and cryptodira. Pleurodiran turtles retract their heads into their shells by bending their necks in the axial lpane whereas cryptodiran turtles lower their necks and retract their heads into their shells. The majority of living turtles are cryptodiran turtles, but there are notable groups of pleurodirans as well. Stupendemys is one such notable member of the pleurodira. the anatomy of this system of hiding its head has been described in many papers, notably in Scheyer and Sanchez-Villagra 2007 and the slightly older de Broin, et al. 1993. The initial description of the turtle is not online; however, the description of the second species named, Stupendemys souzai, is available online. This is a good reference for not only the anatomy of the genus, but also general turtle anatomy, as modern knowledge of turtles is used to more accurately describe the fossil by the authors Bocquentin and Melo.

Big Turtle, Little Star

There is an awful void of popular knowledge about Stupendemys. The name alone, meaning Astonishing Turtle, would seem to require more coverage than the turtle has received on the internet, in documentaries, or in movies and television. The giant turtle is shown some love in the amateur video sphere as more than one fossil enthusiast has dedicated some time to describing the turtle and sharing illustrations and other recreations in their videos. The most informational of these videos was posted by a user from the United Kingdom known as The Sarcastic Raptor who posts videos about games as often as videos about extinct animals. Enjoy his description and the images he has found for his video below.

Fresh Prince of Turtles

Freshwater turtles are capable of growing to large sizes in some genera if allowed to continue to growing for along time and not killed at young ages. The past is littered with the remains of giant turtles, both freshwater and marine. One of the largest genera of these freshwater turtles was the turtle Stupendemys. Consisting of two acknowledges species (S. geographicus Wood 1976 and S. souzai Bocquentin & Melo, 2006), Stupendemys carapaces averaged 1.80 m (5.9 ft) long with the largest individual approaching an estimated 3.3 m (11 ft) long carapace; an estimate that makes this turtle larger than the largest marine turtle Archelon. These two species are of similar size with S. souzai being described as smaller and have both been recovered from South America; S. geographicus from Venezuela and S. souzai from Brazil. Thought to be an enormous grazer and terrible swimmer, it has been hypothesized that Stupendemys used its weight to control its buoyancy and keep it grounded in the vegetation of the calm slow moving riverbeds it would have inhabited. There are hypotheses that the habitat of the turtle may have lacked significant predators, allowing the animal to reach the sizes that it did, but regardless of the validity of these hypotheses, this was an enormous turtle as we can see below with this replica carapace:

Unattributed photo found online

Giants of the Lost World

Donald Prothero has taught multiple generations of paleontologists, biologists, and science enthusiasts in his 35 year career. He has written more papers and books than I can count and I, along with many others in the field, have read a great deal of his work over the years and know his contributions to the field quite well. I was recently offered the opportunity to read and review his latest book, Giants of the Lost World: Dinosaurs and Other Extinct Monsters of South America (Smithsonian Books, 2016) and I am quite honored to share my impressions of the book with all of you here today.

I am a big fan of art and figures in scientific books and so the first thing that I do with any book is check out the cover and thumb through the illustrations. This book, from the artistic standpoint, does not disappoint the modern reader. Images are clearly captioned and placed in appropriate places in coordination with the narration of the book. Photographs, reconstructions, and supporting graphics (phylogenetic trees, maps, etc.) are all black and white, save 16 color plates which also include computer generated models of some taxa. These illustrations are high quality, modern scientific interpretations of extinct animals which, for readers that enjoy visuals when they read scientific books, enhance the reader's understanding of the worlds that are being discussed in this book. What's more, the book's sources for illustrations are taken, often, from the internet; an innovative approach to sourcing illustrations.

I say worlds in reference to the illustrations rather than "the world" because the narrative of this book is not simply the history of South America, but the history of the lands that South America bordered, touched, and influenced (or has been influenced by) in terms of both flora and fauna. Additionally, global extinctions are discussed at the global scale, as they should be. The treatments of the debate over the modes and scopes of great extinctions like that at the Cretaceous- Paleogene boundary is intense with details of geology, biology, and physics explained that caused specific events to occur as we know them from the fossil record. The well laid foundation of historical events in the book is key to discussions of the flora and fauna of different eras and this history is very well detailed. The language of the history is scientific, but easy to approach for casual science enthusiasts like many of the readers of this blog. However, seasoned scientists can appreciate the professional language and how it is made accessible to not only scientists but also the general public in a manner that is professional and respectful.

The treatment of the animals in this books is likewise respectful and equal across taxa. A paleontologist focusing on fossil rhinoceroses and other mammals for a great portion of his career, one might expect Donald Prothero to be extremely biased toward the mammals that he discusses in the book. It is true that a good portion of the book does discuss mammals, their rise to dominance over the landscape, and the many different stages of their evolution; however, reptiles, dinosaurs, and birds receive considerable attention as well and there are even mentions of amphibians and marine life.

This book is written for a popular audience and discusses and describes extremely high level concepts in a manner that is accessible, fun, and well illustrated. The work is educational but entertaining such that it is an easy read. The chapters can be read as stand alone units but together represent a rich history of a continent, its animals and plants, and the world in which it is a part. This is a book that should be read by professional and casual paleontologists that love mammals, reptiles, birds, and the overall pursuit of more knowledge.

Life Sized Models

A model of Nqwebasaurus existing somewhere in the world is inevitable. The fact that it is in South Africa should be of very little surprise given that the fossils were discovered there. This model pre-dates the knowledge of the ubiquity of feathers and as such is quite a basic scaly dinosaur, but feathers in theropods this basal have not been ruled out as entirely possible as yet. The anatomy described in the papers we read this week is evident in this model. The digits of the manus appear to be quite exaggerated; however, in looking at the fossil we can see that the digits are fairly elongated. Impressively, the opposable digit is very well modeled here, making the forelimb look that much more terrifying. Possessing feathers or not possessing them, the models and illustrations we have seen this week of Nqwebasaurus are all equally interesting and make this basal coelurosaurian look quite dynamic.

Photo by William De Klerk

Famous Tongue Click

Nqwebasaurus is the only dinosaur with a name that includes a tongue click. It is also the only basal coelurosaurian from South Africa and one of the earliest theropods found in the entirety of the continent. These two reasons for popularity have fueled inquiry and interest in Nqwebasaurus but its name is difficult to say and spell, which has stalled some of the general popularity that the dinosaur has been subjected to or affected by (depending on your stance). The strange general popularity lack for this dinosaur even makes the illustrations of the dinosaur somewhat rare. There are interesting facts that are less well known about the dinosaur that could make it even more interesting or popular with the general (non-scientist) populace. One of these is the lesser stressed identification of possible gastroliths in the abdomen of the dinosaur, pointing to an herbivorous diet. Gastroliths are stones that are smoothed by mechanical manipulation in a muscular organ such as a gizzard or crop. These stones are used to grind tough fibrous foods so that an animal does not need to chew and rechew and chew its food again to break down the strong vegetation to get at the nutrients contained within.

Opposable Thumbs

If one remembers the cartoon that was shared on Sunday there was mention of opposed digits and climbing abilities prescribed to Nqwebasaurus. There was not much embellishment in those statements; there is always some in developing hypotheses. The forelimb and hand of Nqwebasaurus are unique in that they do appear to have been well-adapted to not only climbing, but also partial opposition of the digits allowing for a certain amount of grasping. This amount of grasping has not been quantified, though the partial nature of the opposition has been stressed in the initial description and subsequent publications. When looking at the fossil it is not difficult to understand why the degree of movement of the manus and its digit's movements is not entirely explored, modeled, or simply known at this time:

The Literate Dinosaur

Nqwebasaurus is well read and well read about. Not many basal coelurosaurs have been found and of those that have, only one comes from South Africa. Officially an ornithomimosaur, Nqwebasaurus has been discussed in theropod, coelurosaurian, and ornithimimid contexts since its discovery and description in 2000. The paper announcing that discovery is a paper worth reading and the author list is little bit like an all star lineup for paleontology. The paper appeared in the Journal of Vertebrate Paleontology and is authored by De Klerk, Forster, Sampson, Chinsamy, and Ross (if you are a more casual reader these names are less impressive than if you are familiar with their work, but it is impressive, I promise). There has been extensive examination of the fossils and subsequent discoveries since the initial description such as a description of a second set of remains in 2009 by Forster, et al. that compares these remains with those of the holotype of Nqwebasaurus. They determine that these belong to a second theropod inhabiting the same area, which is equally intriguing despite not representing another individual of Nqwebasaurus. Placing animals outside of Nqwebasaurus is not the only thing that has been done to the genus since its description though. In 2012 Choiniere, et al. re-examined the holotype material and further excavated the matrix of the fossil. They found more material and were able to better describe Nqwebasaurus which included a more complete and more soundly supported phylogenetic assignment to the position we now know for Nqwebasaurus. This paper is so far the most important description or redescription of the animal and its place in the family tree.

Morning of the Nqwebasaurus

Cartoons used to be a normal way to start weekend mornings and those of us old enough to remember certainly miss them. Kids that love dinosaurs should start today with a cartoon featuring a talking Nqwebasaurus. While watching this short, they can read the facts presented on Prehistoric Wildlife or a more abbreviated version on Enchanted Learning. Adults can enjoy these also, of course.

Early Bird Mimics

There are a lot of dinosaur names that are hard to pronounce or spell and one of the absolute most difficult is the basal ornithomimosaur Nqwebasaurus thwazi. Recovered from South Africa's Kirkwood Formation of the Early Jurassic-Late Cretaceous, Nqwebasaurus is named after the formation itself, which is called Nqweba in Xhosa, the local group of Bantu people living nearest the formation where the skeleton was recovered from. As the earliest coelurosaur of Gondwana, it is remarkable in many ways despite looking somewhat plain:

©IJReid

Pretty Whales

Basilosaurus is a favorite for reconstructions as well as re-creations and interpretations in paleoart (there is currently a lot of talk about the differences on Twitter). There are many different illustrations of Basilosaurus that have many different body types and rather than discuss the merit of one over another today, I feel that is is much more productive to instead share a few and let the readers come to their own conclusions on what is the most whale-like interpretation or the least, or even the most interesting regardless of its whale-like attributes. Please enjoy these interpretations of Basilosaurus and later today I will post about the new animal for this week. Additionally, I am in the process of finishing up a book that will be my pleasure to introduce and briefly discuss with everyone.

© Pavel.Riha.CB CC BY-SA 3.0

Unattributed

©Nobu Tamura CC BY-SA 3.0

Popular Whales

The most common avenue for popularity for any fossil animal outside of documentaries in current society is through video games. The hopes and dreams of many scientifically minded folks always want the number one area to be the museum but with so much electronic distribution of information these days the museum has fallen behind slightly but knowledge about animals. Curiosity generated through any venue or modality is extremely important in moving the research, future knowledge, and continued popularity of fossil animals higher in the public and scientific conscience. Seeing Basilosaurus featuring prominently in video games is somewhat intriguing to me. Its portrayal in Ark: Survival Evolved is as a highly detailed creature that lives within the underwater portion of this game's world. There are a number of videos for the whale, but this one made by Alexis Arcade, despite some interesting pronunciations is directed with vigor and excitement, and that is all I really ask for as long as the information, which in this case is that which is provided by the game creators, is accurately relayed to our audience. Anyone interested in a very long video with the animal swimming, fighting, and eating in game should check out Jimbob Soss' video here (38:00 is where the Basilosaurus comes into the video).

Stereotypical Papers

One of the most highly anticipated discoveries in the whale research is the purpose of vestigial hindlimbs and, heavily coinciding with that topic, when the hindlimbs became entirely vestigial. I bring this up because two and a half (one paper being a description of the entire skeleton and mentioning the limbs and lack thereof) of the first five papers that appear in a standard search for Basilosaurus are concerned with the pelvic girdle an the hindlimbs. The evidence of hindlimbs and feet, discussions on the pelvic girdle (hips) themselves, and a treatise on the jaws and "other parts" figure prominently in Basilosaurus research. As an early whale this is not only expected, but also somewhat unimpressive, given that it is expected. The finds themselves are impressive and tell us a lot about the evolution of whales and what makes a whale a whale. Those of the readers interested in the teeth and biting of Basilosaurus are not left out, there is a wonderful paper on bite marks and the forces associated with Basilosaurus bites in a Dorudon (another type of whale) that is available online to read. This paper includes very clear photographs of Dorudon fossils as well as some nice reconstructions of the wounding action.

Movie Whales

It turns out there are quite a few resources for Basilosaurus in the film and movie department. The most famous is the one we are going to watch, but the others will be around for other days in the future, like Thursday when we talk about popular culture references. This is only a small clip from the show, but it is worth finding the whole episode if one can find it.

Belated Facts

Because I was a little slow this weekend (I was doing a lot of reading mixed with a lot of not being online and hanging out with my wife) I am posting the facts about Basilosaurus on a Monday morning. There are many websites and a number of videos that share facts about Basilosaurus because it is an animal of interest to many people, scientists and non-scientists alike. Websites include About, Prehistoric Wildlife, and even the BBC, just to name a few prominent sites that list facts and discuss this early whale. Since it is Monday and possibly more hectic than a normal day, the videos playing in the background may be easier to get facts from for some today. There are a number of fact presenting videos, but the two below summarizes and discuss the popular knowledge about Basilosaurus equally well:

Angry Whales

The earliest whales walked the shorelines of swamps and oceans. The earliest large fully aquatic whales that we know of and would recognize as whales were still rather alien appearing in terms of what we think of as whales; however, their largest representatives could be mistaken for nothing else due to their size and mammalian characteristics. The king of these early whales, by name the "king lizard", was Basilosaurus. Consisting of two recognized species, B. cetoides and B. isis, Basilosaurus was a predatory whale measuring up to 18 m (59 ft) long and possessing a skull and feeding apparatus capable of an estimated 20,000 pounds per square inch of force during an extreme bite. Flippers extending 35 cm (14 in) stabilized the whale as it propelled itself through the water with a hypothesized undersized, by modern standards, fluke capable vertical movements possessing either a dorsal fin or ridge along the midline of the tail. This structure is hypothesized currently and its exact size and morphology are unknown presently, though support for both morphologies have been published in the past.

Basilosaurus on display in the Life in the Ancient Seas exhibit, 1989. Photo by Chip Clark, Smithsonian Institution

Skinny or Beefy Neck

© Øyvind M. Padron

The neck of Piatnitzkysaurus is of the least concern to most researchers and illustrators. However, in searching for impressive artistic interpretations today I noticed that there was an interesting range of neck morphologies depicted for this dinosaur. Ranging from extremely gracile and snake-like to highly muscular and robust, the illustrations of the neck in Piatnitzkysaurus set the tone for the remainder of the illustration. A dinosaur with a gracile neck is most often portrayed as possessing a gracile frame, hindlimbs, tail, and forelimbs. Typically these smaller necks also have lighter heads resting at their rostral ends; a giant beefy head would be difficult or impossible to support on a gracile neck. Conversely, dinosaur illustrations with extremely large necks tend to have robust and muscular bodies and heads to realistically portray proportional animals. The difference between gracile and robust is not just the overall look of the animal that is reproduced of course. Predatory dinosaurs with smaller heads and more streamlined bodies are most often portrayed as speedy smaller animals capable of stealing eggs and chasing down faster prey items. These tend to influence the dinosaur movie chase scenes that involve acrobatics and high speed escapes. Robust heads and bodies that appear stronger are more common with dinosaurs that break through walls or crush cars in movies in order to get at their prey. Hollywood, in this respect, is not entirely incorrect; nor should they be considering that paleontologists have been consulting for these kinds of movies for a long time. The difference between robust and gracile versions of Piatnitzkysaurus has not been settled and interpretations on both ends of the range are still produced. The anatomically correct version of the animal is somewhere between the two, though examples of both probably existed within the variation of the animal in its history as a genus and neither should be entirely discounted without definitive proof of a body size.

©Rodrigo Henrique Gomes

Long Drawn Out Dig

The holotype remains of Piatnitzkysaurus took multiple years to fully excavate from the Patagonian rocks. Dated from approximately 166 to 164 million years ago, the Callovian rocks of the Jurassic were excavated in 1977, 1982, and 1983 to remove the entire holotype specimen. The general resemblance of this dinosaur to Allosaurus was first noted by Michael Benton upon surveying the strong arms and legs of the predatory dinosaur. Many of the illustrations are very much inspired by this resemblance in addition to the actual anatomy of Piatnitzkysaurus, as can be seen here.

©Nobu Tamura