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Altispinax: The Mysterious Meat-Eater of Early Cretaceous England

Introduction

There are many dinosaur species which have been identified based upon very scant remains, and this article concerns one of them: a meat-eating dinosaur named Altispinax dunkeri. If you’ve never heard of this animal before, you’re not alone. It’s not exactly a name that readily springs to mind when one thinks of dinosaurs. After all, Altispinax is only known from three vertebrae that were found back in the 1850s, and nothing else has been found since then which can be positively attributed to this animal. Other isolated fossils have been found here and there, but there’s really no way to tell if all of these isolated bones, teeth, and claws all belong to the same creature. Broadly, we are reasonably sure that it was a meat-eating dinosaur, but aside from that, there’s not much else to go on.

Discovery and Description

In 1855, a British lawyer and amateur fossil hunter named Samuel H. Beckles (April 12, 1814 – September 4, 1890) was prospecting for fossils in southeastern England. Not far from the site of the Hastings battlefield, he discovered a series of three vertebrae with unusually tall dorsal spines, measuring about 14 inches (35 centimeters) from the top of the centrum disk to the top of the spine (1).

The holotype specimen of Altispinax (collections ID code: BMNH R1828, or maybe it’s NHMUK R1828). Image from “Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia” by Darren Naish and David M. Martill (2007). Journal of the Geological Society, volume 164. Page 503, Figure #7 (Pages 493-510).
https://darrennaish.files.wordpress.com/2013/01/naish-martill-2007-gsl-british-dinosaurs-pt-i-saurischia.pdf.

The following year in 1856, the famous British paleontologist Sir Richard Owen wrote a description of this fossil in which he ascribed it to the genus Megalosaurus, which had been classified about thirty years earlier. The description of the fossil reads as follows:

“Three anterior dorsal vertebrae: p, parapophysis, or lower transverse process: t, accessory tubercle contributing some attachment to the head of the rib: d, diapophyses, or upper transverse process, fractured, which gave attachment to the tubercle of the rib: b, oblique buttress extending from the parapophysis to the diapophysis, and contributing to the support of the neural platform : z, the prozygapophysis, z’, the zygapophysis, forming the ends of the neural platform and articulating the neural arches of the vertebrae with each other, ns, the neural spine of the foremost of these vertebras, ns’, the neural spine of the second vertebra; it expands at its extremity, overhangs the anterior shorter spine, and developes a strong bony plate from its back part which fixes it to n”, the similarly developed and modified spine of the third vertebra. The extraordinary size and strength of the spines of these anterior dorsal vertebrae, indicate the great force with which the head and jaws of the Megalosaurus must have been used” (2).

This is a drawing (yes, that’s a drawing, not a photograph) of the vertebrae, made by Joseph Dinkel which accompanies Owen’s article. Public domain image, Wikimedia Commons.
https://commons.wikimedia.org/wiki/File:Becklespinax.jpg.

The fossil vertebrae were found in the rocks of the “Wealden Supergroup”. This is a multi-layer series of geological formations consisting largely of sandstones and mudstones which are spread throughout much of southeastern England and date to the early Cretaceous Period. This geological super-formation is divided into several sub-units called “groups”, and each of these groups are divided further into “formations”. These include the “Hastings Beds Group” (dated from the Berriasian to Valanginian Stages of the early Cretaceous Period), the “Weald Clay Group” (Hauterivian and Barremian, possibly extending into the Aptian), and the “Wealden Group” (Barremian to Aptian) (3).

The vertebrae fossils were found in the rock layers of the “Hastings Beds Group”. Specifically, the fossils were found in the Wadhurst Clay Formation of East Sussex. This formation dates to the Valanginian Stage of the early Cretaceous Period, and even more specifically to the early to middle Valanginian. This would place the vertebrae’s date range at about 138-135 million years ago. During this same time, iconic early Cretaceous dinosaurs such as Iguanodon and Baryonyx had not yet evolved. Instead, Altispinax shared its habitat with the armored ankylosaur Hylaeosaurus, the sauropods Pelorosaurus and Xenoposeidon, and the small ornithopods Valdosaurus and Echinodon. Isolated fossils of Iguanodon-like animals have also been found here, as well as fossils of frogs, crocodiles, and small mammals. Since no other large carnivorous dinosaurs are known from this place and time, Altispinax might have been the top predator within its environment (4).

Discussion

Altispinax is not a well-known dinosaur, but even though it is not a major star in dinosaur paleontology circles and it is not a creature well-known to the general public, largely due to the fact that the only evidence for it comes from a single fragmentary specimen found nearly 170 years ago, Altispinax has received a surprisingly high amount of academic attention. Every now and then, professional paleontologists dabble in Altispinax speculation, analyzing these three vertebrae for the umpteenth time and pondering where exactly it fits in the theropod family tree.

Unsurprisingly, because these finds are 170 years old and are known only from fragmentary remains, Altispinax’s position in the dinosaur family tree has been regularly shuffled around because nobody, it seems, can make up their minds about what sort of animal it was. When it was first discovered, Sir Richard Owen believed that these spines belonged to Megalosaurus bucklandi, one of the first dinosaurs to be discovered and named. During the 19th and 20th Centuries, the name “Megalosaurus” was considered a “wastebasket taxon” – any remains of a meat-eating dinosaur which could not be positively identified were classified as Megalosaurus. As a result, this genus name acquired a long list of species names. Indeed, early reconstructions of Megalosaurus incorporated these three tall-spined vertebrae into its anatomy. For example, the British sculptor Benjamin Waterhouse Hawkins created a statue of Megalosaurus for Crystal Palace Park in which it possessed a bison-like hump over its shoulders.

Middle 19th Century reconstruction of Megalosaurus by Benjamin Waterhouse Hawkins, on display in Crystal Palace Park. Note the prominent bison-like hump over the shoulder. Photo by C. G. P. Gray (2007). Creative Commons Attribution 3.0 Unported license.
https://commons.wikimedia.org/wiki/File:London_-Crystal_Palace-_Victorian_Dinosaurs_1.jpg
.

When Megalosaurus’ appearance was further refined during the late 1800s into a definitely dinosaurian appearance, but in an incorrect upright pose as championed by people such as Joseph Leidy and Louis Dollo, this reconstruction again showed Megalosaurus with tall neural spines, forming a low ridge running down the middle of its back.

Late 19th Century reconstruction of Megalosaurus by Christian Von Meyer, in which the animal is portrayed with tall neural spines on its dorsal and sacral vertebrae. Illustration from Extinct Monsters, 5th Edition, by Reverend Henry N. Hutchinson. London: Chapman and Hall, 1897. Page 78. https://archive.org/details/extinctmonsters00hutciala.

In 1923, the famed German paleontologist Friedrich Von Huene took a look at the three spines from southern England, as well as a single tooth that was discovered in Germany which had been given the name Megalosaurus dunkeri. Von Huene lumped these fossils together in the belief that they both came from the same species, and determined that they did not, in fact, come from Megalosaurus. So he gave these spines and the solitary tooth a new name – Altispinax dunkeri, “Dunker’s high-spined ruler” (5).

Later researchers justifiably felt that Von Huene had jumped the gun by assuming that the tooth from Germany and the vertebrae from England belonged to the same species. Furthermore, new species had been discovered since then. In the late 1940s in the United States, another large theropod was discovered with a prominent ridge running down its back which looked remarkably similar to the three vertebrae found in England. In 1950, the American dinosaur was named Acrocanthosaurus atokensis, and other paleontologists began to wonder if the mysterious vertebrae found in England came from a similar animal. In the 1980s, paleontologist and paleo-artist Gregory Paul felt that the three vertebrae represented a European version of Acrocanthosaurus, so he gave it a provisional name of “Acrocanthosaurus altispinax”, though he wasn’t absolutely certain if he was correct. In 1991, George Olshevsky stated that these three spines did not belong to Acrocanthosaurus, but belonged to a new genus, and so it was named Becklespinax altispinax, named in honor of Samuel Beckles who had discovered the vertebrae in 1855 (6).

Skeleton of Acrocanthosaurus on display at the North Carolina Museum of Natural Sciences. Photo by Famille Wielosz-Caron (August 4, 2007). Creative Commons Attribution 2.0 Generic license.

Illustration of Becklespinax made by the paleo-artist James Robins in the magazine Dinosaurs!, issue #46, page 1084. Durham: Atlas Editions Partworks, Inc., 1994.

The name Becklespinax remained in place for the next twenty-five years until 2016 when it was challenged by Michael Maisch. He said that since Friedrich Von Huene recognized this animal as a distinct species back in 1923, this pre-dated George Olshevsky’s recognition of this animal as a distinct species in 1991. Therefore, the name Altispinax held priority over the more recent name Becklespinax, and therefore the name Becklespinax should be discarded. So in 2016, the old name of Altispinax dunkeri was resurrected (7).

Just as its name has been altered over the years, Altispinax’s phylogeny has also changed. In 1856, Sir Richard Owen classified this animal as a megalosaur. Then, following the discovery of Acrocanthosaurus in the late 1940s, it was believed to be an allosaur (later, Acrocanthosaurus was re-classified as a carcharodontosaurid). In 1991, George Olshevsky classified Altispinax once again as a megalosaur, specifically in the family Eustreptospondylidae. By the late 1990s, it was re-classified again as being more closely-related to a Middle Jurassic allosaur from China called Sinraptor, and was therefore placed in the allosauroidean family Metriacanthosauridae, which includes Metriacanthosaurus, Sinraptor, and Yangchuanosaurus. These species measured 20-25 feet long (which was the same size estimate commonly seen for Altispinax), had curvaceous skulls, and unusually tiny hands with stubby fingers and claws. Allosaurus, with its gigantic hands and meat hook claws, was a rough-and-tumble attacking predator (numerous injuries on fossils prove this), but its cousin Sinraptor did not have such well-developed hands. In 2003, Darren Naish classified this creature as belonging to the more evolutionarily-advanced super-family Allosauroidea, but didn’t hazard placing it into a specific allosaur family (8). In a 2007 article, Darren Naish wrote “Several other tetanurans exhibit a similar pattern of neural arch laminae to Becklespinax, including Condorraptor, Piatnitzkysaurus and Sinraptor, and Sinraptor in particular exhibits a superficially similar morphology: this could mean that Becklespinax is a sinraptorid, but there are no uniquely shared characters that might demonstrate this. For now, pending further discoveries, Becklespinax remains an indeterminate tetanuran of unknown affinities. Its combination of characters, and uniquely expanded neural spines tips, mean that it is a valid, diagnosable taxon” (9).

Skeleton of Sinraptor. Photo by Ian Armstrong (April 17, 2005). Creative Commons Attribution-Share Alike 2.0 Generic license.

Some people wondered if Altispinax might have been a spinosaur or perhaps a carcharodontosaur, since species from both groups have been found in Europe, and especially considering that carcharodontosaurs like Acrocanthosaurus and spinosaurs like Suchomimus and Ichthyovenator were known to have tall neural spines on its vertebrae forming a dorsal ridge. In a 2007 article, back when the animal was still referred to as Becklespinax, Darren Naish wrote “there are no shared derived characters that might unite Becklespinax with either spinosaurids or carcharodontosaurids, and these suggestions can’t be supported”(10). But not long after this was written, Concavenator was discovered in Spain, and old questions were resurrected regarding Altispinax’s position on the theropod family tree. It was determined that Concavenator was a carcharodontosaurid, and there were several noticeable similarities between this animal and Altispinax. Now, many people are once again changing Altispinax’s phylogeny, saying that, yes, it is a carcharodontosaur, but as to whether or not it is, it’s anybody’s guess at this point. Darren Naish re-classified Altispinax as a carcharodontosaurid in 2011, and the science website Palaeos also classifies Altispinax (identified on the web-page as Becklespinax) as a member of the family Carcharodontosauridae (11).

Skeleton of Concavenator, a 20 foot carcharodontosaurid from Spain. Photograph by Santiago Torralba (August 22, 2008). Creative Commons Attribution 2.0 Generic license.

Reconstructed skeleton of Concavenator. (May 1, 2019). Creative Commons Attribution-Share Alike 4.0 International license.

Reconstruction

Well, time to reconstruct what the whole animal may have looked like. It is not yet clear if Altispinax was more closely related to Sinraptor or to Concavenator. Although both of these animals had different features to the structure of their skull, outwardly they would have looked very similar to each other. Both groups of animals also had short arms with small hands, and would have had an allosaur-ish body structure. One wonders if the carcharodontosaurids evolved directly from the metriacanthosaurids like Sinraptor, but this is just a guess on my part.

As for the sail, that was a bit difficult. For years, paleo-artists depicted sail-backed bipedal dinosaurs, like the old renditions of Spinosaurus, with a sail located on the back between the arms and the legs. But the best position for a theropod’s sail is directly over the hips, not between the front limbs and hind limbs as is often seen. Animals that have their sails positioned between the front legs and back legs are quadrupedal animals, like Dimetrodon, and also what we now know Spinosaurus to be like. In a four-legged animal, the body is held off of the ground in an arch, braced by the four legs, and as such the main body has the structural support necessary to carry a sail over the back. However, in a bipedal animal, the hips form a fulcrum which balances the animal. If a bipedal animal had a sail located in a Dimetrodon-like way, positioned between the arms and legs, it would make the animal front-heavy. Therefore, a sail would be better positioned over the hips. Of course, this discounts the possibility that the animal had an unusually long tail to counter-balance the weight in the front. This is with the assumption that Becklespinax’s sail was short, extending only a small distance along the length of its back. It’s possible that it may have had a skeleton similar to Acrocanthosaurus, which had raised dorsal spines running all along its back, from the base of the skull, and extending down to the tip of the tail. However, this cannot be definitively stated to be the case until more of this animal is discovered.

The drawing that you see below was made with No. 2 pencil.

Altispinax dunkeri. © Jason R. Abdale (May 3, 2021).

Source Citations:

  1. Everything Dinosaur. “Remembering Samuel Husbands Beckles (1814-1890)”, by Mike Walley (August 24, 2014).; I Know Dino podcast. Episode 54 – “Becklespinax” (December 8, 2015).
  2. Richard Owen (1856). “Monograph on the fossil Reptilia of the Wealden Formation. Part IV”. Palaeontographical Society Monographs, volume 10.
  3. Darren Naish, “Pneumaticity, the early years: Wealden Supergroup dinosaurs and the hypothesis of saurian pneumaticity”. In Richard T. J. Moody, E. Buffetaut, Darren Naish, and David M. Martill, eds., Geological Society Special Publication No. 343 – Dinosaurs and Other Extinct Saurians: A Historical Perspective. London: The Geological Society, 2010. Pages 229-230.
  4. David B. Weishampel, ‎Peter Dodson, ‎Halszka Osmólska, eds., The Dinosauria, Second Edition. Berkeley: University of California Press, 2004. Page 73; Darren Naish, “Pneumaticity, the early years: Wealden Supergroup dinosaurs and the hypothesis of saurian pneumaticity”. In Richard T. J. Moody, E. Buffetaut, Darren Naish, and David M. Martill, eds., Geological Society Special Publication No. 343 – Dinosaurs and Other Extinct Saurians: A Historical Perspective. London: The Geological Society, 2010. Page 230; Mindat. “Becklespinax”; British Geological Survey. “Wadhurst Clay Formation”; Oladapo Odunayo Akinlotan (October 2015), The Sedimentolody of the Ashdown Formation and the Wadhurst Clay Formation, Southeast England. PhD Thesis, University of Brighton. Page 1.
  5. Smithsonian Magazine. “B is for Becklespinax”, by Riley Black (October 22, 2012).
  6. Smithsonian Magazine. “B is for Becklespinax”, by Riley Black (October 22, 2012).
  7. Michael W. Maisch (2016), “The nomenclatural status of the carnivorous dinosaur genus Altispinax v. Huene, 1923 (Saurischia, Theropoda) from the Lower Cretaceous of England”. Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen, volume 280, issue 2. Pages 215-219.
  8. George Olshevsky (1991). “A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia”. Mesozoic Meanderings, volume 2. Pages 22-23 (1-196); Theropod Database. “Carnosauria”; Dinosaur Mailing List. “Re: Becklespinax” (February 18, 1997); Darren Naish (2003), “A definitive allosauroid (Dinosauria; Theropoda) from the Lower Cretaceous of East Sussex”. Proceedings of the Geologists’ Association, volume 114. Pages 319-326; Darren Naish and David M. Martill (2007), “Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia”. Journal of the Geological Society, volume 164. Pages 502-503, Figure #7 (Pages 493-510).
  9. Tetrapod Zoology. “Of Becklespinax and Valdoraptor”, by Darren Naish (October 2, 2007).
  10. Tetrapod Zoology. “Of Becklespinax and Valdoraptor”, by Darren Naish (October 2, 2007).
  11. Theropod Database. “Carnosauria”; Naish, Darren (2011). “Theropod Dinosaurs”. In Batten, ed., English Wealden Fossils. The Palaeontological Association. Pages 526-559; Smithsonian Magazine. “B is for Becklespinax”, by Riley Black (October 22, 2012); Palaeos. “Theropoda: Avetheropoda: Carcharodontosauridae”.

Bibliography:

Ceratodus: The Iconic Lungfish of the Mesozoic Era

Ceratodus was a genus of prehistoric lungfish which existed on Earth for a surprisingly long time, from the middle of the Triassic Period approximately 227 million years ago to the beginning of the Eocene Epoch of the Tertiary Period about 55 million years ago – a jaw-dropping span of 172 million years! That’s impressive by ANYBODY’S standards!

Lungfish as a whole are a primitive group of fish. They first appeared during the early Devonian Period about 416 million years ago (MYA), and it’s believed that they represent an evolutionary “missing link” between fish and amphibians. The closest relatives of the lungfish are the coelacanths, meaning “hollow spines”. That’s not surprising, considering that both lungfish and coelacanths have prehistoric origins as well as that both groups are classified as “lobe-finned fish”.

Lungfish do not have individual teeth like many fish today. Instead, they have four large bone plates (two in its upper jaw, and another two in its lower jaw) that were ridged in texture and crowned with thick triangular projections, and were used for crushing and cracking. Many species of modern lungfish feed on worms, freshwater snails, crustaceans, small fish, and amphibians.

Today, there are only six surviving species of lungfish, and all of them are found in hot tropical environments. With the exception of one species found in the Amazon Jungle and another species found in northern Australia, the remaining lungfish species are found in Africa.

  1. The South American Lungfish (Lepidosiren paradoxa), found in the Amazon River.
  2. The Marbled Lungfish (Protopterus aethiopicus), which is found throughout much of eastern and central Africa.
  3. The Gilled Lungfish (Protopterus amphibius), which is also found in eastern Africa.
  4. The West African Lungfish (Protopterus annectens), which is found, not surprisingly, in western Africa.
  5. The Spotted Lungfish (Protopterus dolloi), which inhabits the Congo Jungle of central Africa.
  6. The Australian Lungfish, also called the Queensland Lungfish (Neoceratodus forsteri), found in northeastern Australia. Of all of the extant lungfish species, this one is believed to be the most primitive.

Special attention must be given to the Australian Lungfish (Neoceratodus forsteri), for not only is this species regarded as the most archaic of all of the extant lungfish, but it was once believed to be the sole surviving member of the prehistoric lungfish genus Ceratodus alive in modern times.

Skeleton of Neoceratodus forsteri. From Günther, Albert. “Description of Ceratodus, a Genus of Ganoid Fishes, Recently Discovered in Rivers of Queensland, Australia”. Philosophical Transactions of the Royal Society of London, volume 161 (1871). Plate XXX. https://www.jstor.org/stable/pdf/109041.pdf.

The lower jaw of Neoceratodus forsteri, seen from above. From Krefft, Gerard. “Description of a gigantic amphibian allied to the genus Lepidosiren from the Wide-Bay district, Queensland”. Proceedings of the Zoological Society, volume 16 (April 28, 1870). Page 222. https://ia800405.us.archive.org/16/items/biostor-107043/biostor-107043.pdf.

The genus Ceratodus was established in 1837 by the famed Swiss ichthyologist Louis Agassiz based upon teeth which were found in European rock layers dated to the Triassic and Jurassic Periods. Most Ceratodus fossils that are found consist of isolated tooth plates, and different species have been named based largely upon difference in tooth morphology. Twenty-two species of Ceratodus have been named since the genus was first described in 1837. For a long time, Ceratodus was what is known as a “waste basket taxon” – all North American lungfish fossils were ascribed to this genus, regardless of how different they were from each other. Recently, a careful re-examination of lungfish fossils have revealed that these animals are remarkably different from each other and may constitute numerous genera, not just one. If that’s the case, then the overall lifespan of Ceratodus as a genus may be dramatically shorter than was previously supposed (Günther, Albert. “Description of Ceratodus, a Genus of Ganoid Fishes, Recently Discovered in Rivers of Queensland, Australia”. Philosophical Transactions of the Royal Society of London, volume 161 (1871). Page 512).

File:Ceratodus.jpg

Ceratodus, painted by Heinrich Harder. From Animals of the Prehistoric World (1916). Public domain image, Wikimedia Commons. https://commons.wikimedia.org/wiki/File:Ceratodus.jpg.

Ceratodus’ length varied depending on the species. Most sources which I have seen give an average length of 3 feet long. However, one species of Ceratodus may have reached truly gigantic proportions, possibly reaching 10 to 12 feet long. This estimate is based upon a single bone plate, which is the largest-known of any lungfish. The tooth plate was found in central Nebraska in rocks dated to the Miocene or Pliocene Epochs of the Tertiary Period. Shimada and Kirkland hypothesized that the tooth had been carried into central Nebraska by river from older rock layers that were located further to the west within Wyoming, in rocks dated to either the late Jurassic or early Cretaceous Periods. However, the tooth isn’t as banged up as you would expect from such a long journey. It’s possible that the tooth is endemic to central Nebraska, and if that is the case, 1) Ceratodus was alive in North America for a much longer geologic time span than previously supposed, or 2) This species is mis-identified and belongs to a new un-described genus of giant lungfish which lived in central North America about 5 million years ago, or 3) This was a species which happened to have unusually large teeth within its jaws, and the overall length of the animal was much smaller than the 4 meter estimate given by Shimada and Kirkland. Unfortunately, only one tooth plate has been discovered. Until more specimens are found, everything that we have to say about this specimen needs to be taken with a great degree of skepticism. (Kenshu Shimada and James I. Kirkland, “A Mysterious King-Sized Mesozoic Lungfish from North America”. Transactions of the Kansas Academy of Science, volume 114, issue 1 (2011). Pages 135-141. https://www.researchgate.net/publication/261964060_A_Mysterious_King-Sized_Mesozoic_Lungfish_from_North_America).

For the artwork accompanying this article, I decided to change up my style. For this drawing, I chose to evoke the whimsical style of the paleo-art of Patricia Bujard. If you don’t know who Patricia Bujard is, then I highly recommend that you check out her work. She is a children’s author and illustrator with a love for prehistoric life, and I find her artwork adorable. There aren’t too many people who can make an Allosaurus “cute”, but dag-nabbit, she somehow manages to pull it off. You can see her artwork on her WordPress page, Pete’s Paleo Petshop. My own drawing, which you can see below, was made with an ordinary Crayola black marker.

Ceratodus © Jason R. Abdale. February 9, 2021.

Keep your pencils sharp, and in this case, also keep your markers properly stored so they don’t dry out.

Caturus

This is Caturus, a prehistoric fish which swam in the oceans during the Mesozoic Era. Fossils of this saltwater fish have been found in North America, Europe, northern Africa, and as far as China within rocks spanning from the beginning of the Triassic Period about 250 million years ago (MYA) up to the middle of the Cretaceous Period, about 100 MYA. However, most Caturus fossils have been found in Europe within rock layers dated to the middle and late Jurassic Period, about 170-150 MYA.

Despite a superficial resemblance to a salmon, Caturus was actually more closely related to a bowfin (Amia calva), which is a rather primitive ray-finned fish.

So far, paleontologists have identified fourteen species of Caturus. The largest species, Caturus furcatus, which lived in the Tethys Sea (the shallow sea that covered much of Europe during the middle Jurassic to the middle Cretaceous Period), reached three feet long; other species were much smaller. One species, Caturus dartoni, is known from North America in rocks dated to the middle Jurassic Period, about 165 MYA. Only two skeletons of this particular species have been found, the largest measuring 15 inches long.

Caturus. © Jason R. Abdale. September 5, 2020.

This drawing was made on printer paper with No. 2 pencil, No. 3 pencil, Crayola colored pencils, Prismacolor colored pencils, and Artist’s Loft colored pencils.

Champsosaurus: The Croc-Lizard of the Cretaceous

When most people hear the words “aquatic reptile”, they usually think of two things: turtles and crocodilians. Some clever people might mention sea snakes, and others might mention marine iguanas. Those who are keen on impressing you may bring up some obscure species like the water monitor, the basilisk lizard, and other species of snakes which venture into water.

In prehistoric times, the list of options that you could choose from was much more expansive. In fact, there were animals around then which aren’t around today which fit into this category. One such group of prehistoric water-going reptiles was known as the “choristoderans” (pronounced as Kore-RISS-toe-DEER-rans).

The choristoderans were a group of semi-aquatic reptiles which lived during the Mesozoic Era. Although not as well-known as other non-dinosaurian reptiles of the Mesozoic such as pterosaurs and ichthyosaurs, they nevertheless shared their environments with dinosaurs for a span of approximately 110 million years and even survived the dinosaur extinction. Choristoderans first appeared during the middle of the Jurassic Period about 175 MYA. The oldest-known genus which is recognizably a choristoderan was Cteniogenys, which measured just one and a half feet long and was very lizard-like in appearance. In life, it probably resembled a small monitor lizard and it likely filled a similar ecological niche. However, the heyday for the choristoderans occurred during the early Cretaceous Period from about 144 to 100 MYA, after which they went into decline. They were fortunate to survive the K-T Extinction, but they were always second fiddle to their crocodile neighbors. Most of the surviving species went extinct about 50 MYA, with the remainder just barely hanging on. The last of the choristoderans completely went extinct around 20 MYA.

The choristoderans belonged to a group of vertebrates called the “diapsids”, meaning that they had two holes in their skull behind each eye socket. Lizards, snakes, crocodilians, pterosaurs, dinosaurs, and birds are all classified as diapsids.

At first glance, choristoderans might be mistaken for crocodiles. However, despite their crocodile-like appearance, they are more closely related to lizards than to crocodiles, at least according to a study made by Mike Lee in 2013 (“Turtle origins: insights from phylogenetic retrofitting and molecular scaffolds”). Their placement in the reptile tree is primarily based upon the structure and arrangement of their ear bones, which is more advanced than those seen in lizards but not as advanced as those seen in crocodilians and birds. Also, the skulls of choristoderans are structurally more lizard-like than crocodilian.

The order Choristodera is divided into four families: Champsosauridae, Hyphalosauridae, Monjurosuchidae, and Simoedosauridae. The more primitive the species, the more lizard-like it is in form. The more derived, then the more crocodilian it is in appearance. The most primitive choristoderans were the monjurosuchids, which looked similar to the modern-day Water Monitor Lizard (Varanus salvator). Even at this early stage in their development, there is fossil evidence that some species like Monjurosuchus possessed webbed fingers and toes. Already, they were adapted to living a semi-aquatic lifestyle.


Skeleton of Monjurosuchus splendens, a primitive choristoderan from China. Photograph by Jonathan Chen (June 13, 2019). Creative Commons Attribution-Share Alike 4.0 International license. https://commons.wikimedia.org/wiki/File:Monjurosuchus-Beijing_Museum_of_Natural_History.jpg

Even more advanced were the hyphalosaurids, which bear a remarkable resemblance to the earlier nothosaurs and thalattosaurs of the Triassic Period. Form tends to follow function in evolution, and these creatures almost certainly led a similar lifestyle. The act of species from completely different groups evolving into more-or-less the same shape is called “convergent evolution”.

The champsosaurids and the simoedosaurids are the most crocodile-like in appearance, and together they form the super-family Neochoristodera. Like crocodiles, these creatures were almost certainly living as shallow-water ambush predators, fitted with long slender jaws lined with small conical teeth. Like modern-day gharials, they may have been primarily or even exclusively fish-eaters.

Probably the most famous choristoderan genus was Champsosaurus (pronounced as CHAMP-so-SORE-us). It first appeared about 90 MYA during the Turonian Stage of the late Cretaceous Period, persisted through the K-T Extinction, and finally went extinct during the Paleocene Epoch of the Tertiary Period about 56 MYA. Impressive. Most genera don’t last that long.

Champsosaurus was named by the famed paleontologist Edward D. Cope in the year 1877. Despite not having an easily-recognizable name (most members of the general public have likely never heard of it), it has been rigorously studied by paleontologists ever since then. For example, three academic articles were published about it just in the year 2010, and another article was recently published in April 2020. So, from an academic standpoint, interest in this animal has been pretty consistent.

There are seven species which have been ascribed to the genus Champsosaurus. Most of them measured 5 feet long or thereabouts, but the largest, which was appropriately named Champsosaurus gigas, reached 10 feet long. Most Champsosaurus fossils have been found in south-central Canada and the north-central United States within rocks dated to the late Cretaceous Period from 90 to 66 MYA, but a few have also been found in Belgium and northern France in rocks dated to the Tertiary Period.

Champsosaurus skeleton from Montana, USA on display in the Royal Ontario Museum. Photograph by Daderot (November 21, 2011). Public domain image, Wikimedia Commons. https://commons.wikimedia.org/wiki/File:Champsosaurus_sp.,_Montana,_USA,_Late_Cretaceous_-_Royal_Ontario_Museum_-_DSC00088.JPG.

 

Upper jaw of Champsosaurus, above view (left) and underside view (right). The skull’s length measures about 13 inches. Illustration by Samuel W. Williston. From The Osteology of the Reptiles (1925). Public domain image, Wikimedia Commons. https://commons.wikimedia.org/wiki/File:The_Osteology_of_the_Reptiles_p76.png.

Champsosaurus appears to have been able to tolerate both freshwater and saltwater environments. Fossils of a species called Champsosaurus laramiensis have been found in rocks from the Fox Hills Formation, a geological layer which represents a coastal or estuary environment on the edge of the Western Interior Sea. Fossils of mosasaurs and dinosaurs including Tyrannosaurus have also been found in these rocks.

Preserved skin impressions show that, unlike many lizards, choristoderans like Champsosaurus did NOT have overlapping scales. Instead, the skin consisted of tiny non-overlapping scales, with no crocodile-like dorsal scutes, giving it a very smooth-skinned appearance when seen from a distance.

Unlike crocodiles, which have their nostrils on the top of their upper jaw, Champsosaurus had its nostrils on the front tip of its upper jaw. Perhaps they would use their long nose like a snorkel, sticking just the tip out of the water’s surface in order to stay as concealed as possible.

Champsosaurus had a pair of long thin gharial-like jaws lined with tiny conical teeth. Because of its close affinity towards lizards than to crocodiles, it is highly likely that Champsosaurus had lips and a fully enclosed mouth. But that’s just speculation based upon phylogenic relationships to other reptiles. In terms of hard physical evidence, the teeth themselves are quite small, and are inset from the edge of the jawline rather than standing on the rim of the jaw like a crocodile. This suggests that Champsosaurus had lips covering its teeth like a lizard, unlike crocodiles which don’t have lips.

Compared with crocodilians, the eye sockets of choristoderans are positioned much further forwards on the skull, located halfway or two-thirds of the way back from the tip of the snout. This provides more space for jaw muscles, and the temporal fenestrae (the holes in the back of the skull that accommodate the jaw muscles) were very large in proportion with skull size. Champsosaurus, in particular, had very large temporal fenestrae, which indicates that it had strong jaw muscles and could quickly snap its mouth shut within a fraction of a second – an important adaptation if your diet consists primarily of small fish.

Unlike lizards, Champsosaurus might not have had external ears. Analysis of its skull structure shows that Champsosaurus had internal ears, similar to turtles. This is an important adaptation if you are spending much of your life in the water. Therefore, you would not have seen a pair of ear holes on a Champsosaurus head. Instead, there likely just would have been a slight depression (or maybe not even that) on the side of the head marking where the tympanum (the part of the ear that vibrates in order to make a sound) would have been.

If you spend much of your life in the water, walking really isn’t an issue. Therefore, the limbs of choristoderans are not well-developed. In fact, the more “advanced” the species, the weaker its limb bones appear to be. Champsosaurus is no exception to this – its legs are downright puny in comparison with its body. The bones that make up the arms and legs are short and stumpy, and the hands and feet are small, although the feet are noticeably bigger than the hands. The fingers and toes are thin and end with very tiny claws. This was an animal that would have had a hard time pushing itself onto land. However, there is some evidence that females had more robustly-built limbs than the males due to the need to haul themselves onto land in order to lay their eggs.

The tail of Champsosaurus was flattened, and looked more like that of a crocodile or even a mosasaur than to a lizard. Even so, this animal was definitely not a power-swimmer. If it was, then one would expect the tail to be both longer and broader. Instead, the tail seems to be peculiarly under-developed. Keep in mind, though, that this was likely not an animal that was actively chasing after its prey. If all it was doing was hunkering down on the bottom of a lake or river and waiting motionless for fish to carelessly swim by, then it doesn’t need a well-built tail that’s designed for plowing through the water.

Skeleton of Champsosaurus laramiensis. From “The Osteology of Champsosaurus”, by Barnum Brown (1905). Memoirs of the American Museum of Natural History, volume 9, part 1. Public domain image. http://commons.wikimedia.org/wiki/File:Large_williston_champsosaurus.jpg.

Below is a drawing made of Champsosaurus laramiensis drifting about in a murky pond or stream somewhere in Montana during the late Cretaceous Period. This five-foot-long piscivore would have shared this environment with alligators, crocodiles, turtles, large freshwater fish like gars, sturgeons, and bowfins, and of course dinosaurs like Triceratops and Tyrannosaurus. The drawing was made with No.2 pencil on printer paper.

Anyways, keep your pencils sharp.

Evidence of Therizinosaurs in North America during the Late Cretaceous Period

Introduction

For many years, paleontologists have known about the presence of therizinosaurs (formerly classified as segnosaurs) in Asia, especially within what’s now Mongolia and China. However, Asia and North America were linked during a considerable portion of the Cretaceous Period, and this resulted in an interchange of faunas between the two continents, notably ceratopsians, pachycephalosaurs, tyrannosaurs, and maniraptorans. Could therizinosaurs, which had hitherto been exclusively Asian, have lived in North America as well?

A pair of Tarbosaurus attacking a herd of Therizinosaurus somewhere in Mongolia, approximately 80 million years ago. © Gregory S. Paul (1988). Image used with permission.

 

During the early 2000s, that question was answered with a definitive “yes”. Two genera of therizinosaurs have been described from North America, named Falcarius and Nothronychus. Falcarius represents possibly the earliest stage in therizinosaur evolution, dated to the early Cretaceous Period, while Nothronychus is much larger and more advanced and is dated to the middle Cretaceous. The presence of these two creatures clearly shows that therizinosaurs existed in North America, but so far they have only been found in rocks dated to the early and middle parts of the Cretaceous Period. One wonders if therizinosaurs managed to stay in North America right up until the end of the Mesozoic, 66 million years ago. Would they have kept evolving, becoming larger and more advanced? Would they have lived alongside Triceratops and Tyrannosaurus? (1)

It just so happens that there are a few pieces of evidence here and there which suggest that therizinosaurs did survive past the middle Cretaceous within North America, and that they kept living in North America up to the end of the Cretaceous Period.

 

The Evidence

The idea that there were therizinosaurs in late Cretaceous North America was first proposed by the German paleontologist Hans Sues in 1978. Specifically, he was writing about a particular specimen that had been uncovered in the Dinosaur Park Formation, located in Alberta, Canada, in rocks dated to the Campanian Stage of the Cretaceous Period. The specimen in question was a single “frontal” bone, which forms part of the skull. Today, this specimen is in the collections of the Carnegie Museum of Natural History, categorized as “CMN 12355” (NOT 12349 as you’ll sometimes see in internet searches). In his paper, Sues thought that this frontal bone belonged to a “raptor” dinosaur, and listed it as “gen. et sp. indet.”, which is an abbreviated Latin way of saying “genus and species undetermined” (2).

“CMN 12355”: A frontal bone which may belong to a therizinosaur. Left top: ventral view. Right top: dorsal view. Left bottom: lateral view. Right bottom: medial view. © Tracy Ford. Image from Paleofile.com. Used with permission. http://www.paleofile.com/Dinosaurs/Theropods/Segnosaurincertae.asp

 

Saying that this bone belonged to a raptor is understandable, since the dromaeosaurs and the therizinosaurs are related to each other. Both groups are located in a clade called the “maniraptorans”, which includes the ornithomimids, the oviraptorosaurs, the therizinosaurs, and famously, the dromaeosaurs and troodontids – the so-called “raptors” with their famous killing claws.

The second piece of evidence came in the early to mid 1980s. A single bone called an “astragalus”, which forms part of the ankle, was found in the Hell Creek Formation in rocks dated to the very end of the Cretaceous Period. In 1984, the Canadian paleontologist Dale Russell listed this single peculiar find in a long list of specimens uncovered in the Hell Creek Formation during the middle 1980s. However, this particular specimen has never been analyzed or described in a publication exclusively devoted to this bone. It is simply listed as “therizinosaurid indet.”. In 1992, Kenneth Carpenter looked at this bone, and concluded that it actually belonged to Tyrannosaurus, not a therizinosaur (3).

In 1987, the Canadian paleontologist Philip Currie, who is widely acknowledged as the world’s expert on meat-eating dinosaurs, took a second look at the frontal bone which Sues had examined in the late 1970s, and concluded that Hans Sues had made a mistake. It wasn’t a raptor, but was instead a “segnosaur”, which was the way therizinosaurs were called back then. Currie stated that the bone looked similar to the frontal bone of an Asian therizinosaur called Erlikosaurus, and so he reclassified the bone as “cf. Erlikosaurus” (4).

In 1992, Philip Currie did a more thorough examination of possible therizinosaur finds in Canada. He again wrote about the frontal bone which was initially described in 1978, but he also added two more specimens to the discussion table, both of which were housed in the collection of the Royal Tyrell Museum of Paleontology (RTMP). These specimens were given the identification codes “RTMP 81.16.231” (again, Currie classified this specimen as “cf. Erlikosaurus”) and “RTMP 79.15.1” (a “pedal ungual”, or foot claw, which was classified as “cf. therizinosaurid”) (5).

“RTMP 79.15.1”: A foot claw which may belong to a therizinosaur. © Tracy Ford. Image from Paleofile.com. Used with permission. http://www.paleofile.com/Dinosaurs/Theropods/Segnosaurincertae.asp

 

In 2001, Michael Ryan and Anthony Russell conducted their own analysis of North American therizinosaur finds. They confirmed Currie’s claim that the frontal bone found in 1978 did indeed come from a therizinosaur. They also wrote about a neck vertebra found in the Scollard Formation (specimen identification code is “RTMP 86.207.17”), which dates to the very end of the Cretaceous Period, and which they classified as “Therizinosauridae indet.” (6).

Body fossils of therizinosaurs may be rare in North America, but footprints which may belong to therizinosaurs are more abundant. The first footprints were discovered in the 1990s in the Harebell Formation of northwestern Wyoming. According to an article published in 1996, these footprints were unique because they looked like theropod prints except that they had four toes instead of three – unique among theropods, therizinosaurs have four main toes. The authors postulated that the footprints belonged to an animal whose physical remains had not yet been discovered (7).

In 2011, a single therizinosaur footprint was discovered in Denali National Park, Alaska. The rock that the footprint was found in was part of the Cantwell Formation, which spans 80-65 MYA, and the footprint was placed in a layer dated to about 71-69 MYA. Depending upon which source that you read concerning geological dating, this date of 71-69 MYA either marks the boundary between the where the Campanian Stage ends and the Maastrichtian Stage begins, or else it is the earliest phase of the Maastrichtian Stage. In 2012, Anthony R. Fiorillo of the Perot Museum of Nature and Science (located in Dallas, Texas) published an article concerning this peculiar footprint (8). You can see a photo of it here.

In 2013 and 2014, Anthony R. Fiorillo and a team of other researchers returned to the site in Denali National Park and found a total of thirty-one therizinosaur footprints, along with numerous hadrosaur footprints as well. Like the first footprint that had been found in 2011, all of the other footprints were in rock dated to 71-69 MYA. The fact that footprint trackways of both hadrosaurs and therizinosaurs were found together might indicate that these animals traveled together, possibly for mutual protection. An article was published in August 2018 detailing these discoveries (9).

 

Species Identification

As we have seen in the previous section, there is some evidence in the way of footprints and a handful of isolated bones which suggests that therizinosaurs inhabited North America during the late Campanian or early Maastrichtian Stages of the Cretaceous Period. However, is there any way that we can identify which particular genus or species that these fossils belong to?

The subject of identification has been especially contentious concerning the footprints that were found in Wyoming and Alaska. So far, footprints form the majority of finds that are attributed to late Cretaceous therizinosaurs within North America. The problem is that it is difficult to identify a particular genus or species based solely on footprints, unless the shape of the footprint is extremely distinctive. Another problem is that while footprints are abundant, very few body fossils have been found, and none of them are highly diagnostic. Most researchers who examined them determined vaguely that the creature was a therizinosaur, but they couldn’t be more specific than that, with the exception of Philip Currie who proposed that they might belong to Erlikosaurus or a creature very similar to it.

Because it is so difficult to match a footprint with a particular animal, paleontologists often ascribe footprints their own genus and species names. This is what is referred to as an “ichnogenus”, which is a genus of animal known only from trace fossils, such as footprints, rather than actual physical body fossils.

In the 1996 article which discussed the unusual footprints found in Wyoming, the footprints were ascribed to the ichnogenus Exallopus (pronounced as Ex-ALLO-pus, meaning “from different foot” due to its unusual shape) and its species name was given as Exallopus lovei. The type specimen is identified as “DMNH 5989”, and it was identified as a coelurosaur. According to the website Fossilworks, “Its type locality is Whetstone Creek tracksite, which is in a Maastrichtian terrestrial sandstone in the Harebell Formation of Wyoming” (10). The following year in 1997, the genus name was changed from Exallopus to Saurexallopus (SORE-ex-ALLO-pus), because the name Exallopus was already taken by a species of marine worm (11). Another species, Saurexallopus zerbsti, was named in a 2003 article. The type specimen is identified as “CUMWC 224.2”. According to Fossilworks, “Its type locality is Zerbst Ranch Tracksite, which is in a Lancian fluvial sandstone/sandstone in the Lance Formation of Wyoming” (12). In 2014, a third species was named called Saurexallopus cordata based upon a single footprint fount in British Columbia, Canada, and dated to the Wapiti Formation of the late Cretaceous Period (13).

While all of the scientific articles concerning Saurexallopus identify it as a theropod, there has been some dispute as to what particular type of theropod it is. The original article which was written in 1996 identified it as a coelurosaur. In 2012, Anthony Fiorillo and Thomas Adams identified Saurexallopus as a therizinosaur (14). In an article written in 2015, Saurexallopus was identified as an oviraptorid (15). In an article written in 2018, Saurexallopus was simply identified as a theropod without any specific affinity (16). The website Fossilworks identifies Saurexallopus as a therizinosaur (17).

 

Reconstructing Saurexallopus

During the late 1980s and early 1990s, Philip Currie made comparisons between the various finds in North America with the Asian species Erlikosaurus. According to a phylogenic analysis of therizinosaur genera which was conducted in 2019, Erlikosaurus was closely related to Nothronychus, a therizinosaur which lived in North America during the middle Cretaceous Period. Since Saurexallopus is believed to be physically similar to Erlikosaurus, it is likely that it was genetically related as well, and as such would have been genetically related to Nothronychus. It is therefore quite possible that Erlikosaurus, Nothronychus, and Saurexallopus would have been similar in appearance (18).

Erlikosaurus skull and foot.jpg

Upper jaw and right foot of the Asian therizinosaur Erlikosaurus. Saurexallopus was probably similar in appearance to this genus. Illustration from Rinchen Barsbold and Altangerel Perle (1980) “Segnosauria, a new infraorder of carnivorous dinosaurs”. Acta Palaeontologica Polonica, 25 (2): pages 187-195. https://www.app.pan.pl/article/item/app25-187.html. Creative Commons Attribution License.

 

We can guess that Saurexallopus reached a similar length to Erlikosaurus, measuring about fifteen to twenty feet long (Holtz claims that Erlikosaurus was smaller than other authors do, although his estimate of Nothronychus is in fitting with the size bracket mentioned above) (19). Unlike the eponymous Therizinosaurus, which possessed long scythe-like finger claws (hence its name, which translates to “scythe lizard”), Nothronychus possessed shorter hook-shaped claws, which looked very similar to the stereotypical talons that are seen on carnivorous dinosaurs like Allosaurus and Torvosaurus. These claws were only one-third the size of the claws of Therizinosaurus, but they were well-suited for pulling down branches, for digging (if they could pronate their hands, but that’s a whole other argument), and for smacking the daylights out of any would-be predator. Thomas R. Holtz Jr. has compared therizinosaurs to the large ground sloths of the Cenozoic Era, and the analogy has some merit (20). Saurexallopus and other therizinosaurs likely lived a similar lifestyle and occupied a similar ecological niche, with the possible exception of Falcarius, which may have had a more cursorial lifestyle similar to early coelurosaurs like Ornitholestes.

Based upon their place within the dinosaur family tree, as well as from fossil finds, we are fairly certain that therizinosaurs were feathered. Therefore, it is almost certain that Saurexallopus would have had some form of feather covering as well, although whether it was over the entire body or only partially cannot be determined.

Below is a drawing that I made of Saurexallopus, based upon Erlikosaurus and Nothronychus. The erect mane running down the middle of its neck, back, and tail are just artistic conjecture.

Saurexallopus. © Jason R. Abdale. May 7, 2020.

 

Conclusions

So where does all of this information lead us? So far, there is some evidence which suggests that therizinosaurs were living in Alberta, Canada and Alaska, USA during the late Campanian Stage or early Maastrichtian Stage of the late Cretaceous Period up until about 70 MYA or thereabouts. As such, they would have lived side-by-side with creatures such as Albertosaurus, Edmontosaurus, and Hypacrosaurus. There is only one piece of evidence, a single neck vertebra, which suggests that therizinosaurs existed in North America during the Maastrichtian Stage of the Late Cretaceous. However, no specimens that can be definitely and unquestionably identified as belonging to a therizinosaur have been found in the Hell Creek Formation. Therefore, as far as our current evidence goes, it is unlikely that therizinosaurs lived side-by-side with Triceratops and Tyrannosaurus. However, this may change in the future if more body fossils are discovered.

 

Sources

  1. Utah’s Dino Graveyard; When Dinosaurs Roamed America.
  2. Lindsay Elizabeth Zanno. A Taxonomic and Phylogenetic Reevaluation of Therizinosauria (Dinosauria: Theropoda): Implications for the Evolution of Maniraptora. PhD dissertation, submitted to the University of Utah. December 2008. Page 172.
  3. Lindsay Elizabeth Zanno. A Taxonomic and Phylogenetic Reevaluation of Therizinosauria (Dinosauria: Theropoda): Implications for the Evolution of Maniraptora. PhD dissertation, submitted to the University of Utah. December 2008. Page 172; Dinosaur Mailing List. “Re: Yet even more questions (and I’m sure there’ll be more…)”, by Mickey Mortimer (June 22, 2002). http://dml.cmnh.org/2002Jun/msg00369.html; Theropod Database. “Therizinosauroidea”. http://theropoddatabase.com/Therizinosauroidea.htm.
  4. Lindsay Elizabeth Zanno. A Taxonomic and Phylogenetic Reevaluation of Therizinosauria (Dinosauria: Theropoda): Implications for the Evolution of Maniraptora. PhD dissertation, submitted to the University of Utah. December 2008. Page 172.
  5. Lindsay Elizabeth Zanno. A Taxonomic and Phylogenetic Reevaluation of Therizinosauria (Dinosauria: Theropoda): Implications for the Evolution of Maniraptora. PhD dissertation, submitted to the University of Utah. December 2008. Page 172; Dinosaur Mailing List. “Re: Yet even more questions (and I’m sure there’ll be more…)”, by Mickey Mortimer (June 22, 2002). http://dml.cmnh.org/2002Jun/msg00369.html.
  6. Lindsay Elizabeth Zanno. A Taxonomic and Phylogenetic Reevaluation of Therizinosauria (Dinosauria: Theropoda): Implications for the Evolution of Maniraptora. PhD dissertation, submitted to the University of Utah. December 2008. Page 172.
  7. J. D. Harris, K. R. Johnson, J. Hicks and L. Tauxe (1996). “Four-toed theropod footprints and a paleomagnetic age from the Whetstone Falls Member of the Harebell Formation (Upper Cretaceous: Maastrichtian), northwestern Wyoming”. Cretaceous Research, 17: 381-401.
  8. Anthony R. Fiorello and Thomas L. Adams (2012). “A therizinosaur track from the Lower Cantwell Formation (Upper Cretaceous) of Denali National Park, Alaska”. Palaios, 27: 395-400.
  9. Anthony R. Fiorello and Thomas L. Adams (2012). “A therizinosaur track from the Lower Cantwell Formation (Upper Cretaceous) of Denali National Park, Alaska”. Palaios, 27: 395-400; “The Lower Cantwell Formation and Its Fossils”; “Therizinosaur: prehistoric predator set standard for ‘weird’ in Alaska”; “First North American co-occurrence of Hadrosaur and Therizinosaur tracks found in Alaska”.
  10. Fossilworks. “Saurexallopus lovei”. http://fossilworks.org/bridge.pl?a=taxonInfo&taxon_no=65844.
  11. J. D. Harris, K. R. Johnson, J. Hicks and L. Tauxe (1996). “Four-toed theropod footprints and a paleomagnetic age from the Whetstone Falls Member of the Harebell Formation (Upper Cretaceous: Maastrichtian), northwestern Wyoming”. Cretaceous Research, 17: 381-401; J. D. Harris (1997). “Four-toed theropod footprints and a paleomagnetic age from the Whetstone Falls Member of the Harebell Formation (Upper Cretaceous: Maastrichtian), northwestern Wyoming: a correction”. Cretaceous Research, 18: 139.
  12. Martin G. Lockley, G. Nadon, and Philip J. Currie. (2003). “A diverse dinosaur-bird footprint assemblage from the Lance Formation, Upper Cretaceous, eastern Wyoming; implications for ichnotaxonomy”. Ichnos, 11: 229-249; Fossilworks. “Saurexallopus zerbsti”. http://fossilworks.org/bridge.pl?a=taxonInfo&taxon_no=81011.
  13. R. T. McCrea, L. G. Buckley, A. G. Plint, Philip J. Currie, J. W. Haggart, C. W. Helm, and S. G. Pemberton (2014). “A review of vertebrate track-bearing formations from the Mesozoic and earliest Cenozoic of western Canada with a description of a new theropod ichnospecies and reassignment of an avian ichnogenus”. In Lockley Martin G.; Lucas, Spencer G., eds. New Mexico Museum of Natural History & Science. Bulletin 62: Fossil Footprints of Western North America. Albuquerque: New Mexico Museum of Natural History & Science, 2014. Page 87.
  14. Anthony R. Fiorello and Thomas L. Adams (2012). “A therizinosaur track from the Lower Cantwell Formation (Upper Cretaceous) of Denali National Park, Alaska”. Palaios, 27: 395-400.
  15. R. T. McCrea, D. H. Tanke, L. G. Buckley, M. G. Lockley, J. O. Farlow, L. Xing, N. A. Matthews, C. W. Helm, S. G. Pemberton and B. H. Breithaupt (2015). “Vertebrate ichnopathology: pathologies inferred from dinosaur tracks and trackways from the Mesozoic”. Ichnos, 22 (3–4): 235-260.
  16. Martin Lockley, Gerard Gierlinski, Lidia Adach, Bruce Schumacher, and Ken Cart (2018). “Newly Discovered Tetrapod Ichnotaxa from the Upper Cretaceous Blackhawk Formation, Utah”. In Spencer G. Lucas and Robert M. Sullivan, eds. New Mexico Museum of Natural History and Science. Fossil Record 6, Volume 2: Bulletin 79. Albuquerque: New Mexico Museum of Natural History and Science, 2018. Pages 469-480.
  17. Fossilworks. “Saurexallopus”. http://fossilworks.org/bridge.pl?a=taxonInfo&taxon_no=65843.
  18. Scott Hartman, Mickey Mortimer, William R. Wahl, Dean R. Lomax, Jessica Lippincott, and David M. Lovelace (2019). “A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight”. PeerJ, 7: e7247. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6626525/.
  19. David Lambert, The Dinosaur Data Book (New York: Avon Books, 1990), page 61; Don Lessem and Donald F. Glut, The Dinosaur Society Dinosaur Encyclopedia (New York: Random House, 1993), page 184; Peter Dodson, The Age of Dinosaurs (Lincolnwood: Publications International Ltd., 1993), page 142; Thomas R. Holtz Jr, Dinosaurs: The Most Complete, Up-To-Date Encyclopedia for Dinosaur Lovers of All Ages (New York: Random House, 2007), page 382.
  20. Thomas R. Holtz Jr, Dinosaurs: The Most Complete, Up-To-Date Encyclopedia for Dinosaur Lovers of All Ages (New York: Random House, 2007), page 147.

 

Bibliography

Books:

  • Dodson, Peter. The Age of Dinosaurs. Lincolnwood: Publications International Ltd., 1993.
  • Holtz Jr., Thomas R. Dinosaurs: The Most Complete, Up-To-Date Encyclopedia for Dinosaur Lovers of All Ages. New York: Random House, 2007.
  • Lambert, David. The Dinosaur Data Book. New York: Avon Books, 1990.
  • Lessem, Don; Glut, Donald F. The Dinosaur Society Dinosaur Encyclopedia. New York: Random House, 1993.

Articles:

  • Fiorello Anthony R.; Adams Thomas L. (2012). “A therizinosaur track from the Lower Cantwell Formation (Upper Cretaceous) of Denali National Park, Alaska”. Palaios, 27: 395-400.
  • Fiorillo, Anthony R.; McCarthy, Paul J.; Kobayashi, Yoshitsugu; Tomsich, Carla S.; Tykoski, Ronald S.; Lee, Yuong-Nam; Tanaka, Tomonori; Noto Christopher R. (August 3, 2018). “An unusual association of hadrosaur and therizinosaur tracks within Late Cretaceous rocks of Denali National Park, Alaska”. Scientific Reports, 2018; 8 (1) DOI: 10.1038/s41598-018-30110-8. https://www.nature.com/articles/s41598-018-30110-8.
  • Harris, J. D.; Johnson, K. R.; Hicks, J.; Tauxe; L. (1996). “Four-toed theropod footprints and a paleomagnetic age from the Whetstone Falls Member of the Harebell Formation (Upper Cretaceous: Maastrichtian), northwestern Wyoming”. Cretaceous Research, 17: 381-401.
  • Harris, J. D. (1997). “Four-toed theropod footprints and a paleomagnetic age from the Whetstone Falls Member of the Harebell Formation (Upper Cretaceous: Maastrichtian), northwestern Wyoming: a correction”. Cretaceous Research, 18: 139.
  • Hartman, Scott; Mortimer, Mickey; Wahl, William R.; Lomax, Dean R.; Lippincott, Jessica; Lovelace, David M. (2019). “A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight”. PeerJ, 7: e7247. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6626525/.
  • Lockley, Martin G.; Nadon, G.; Currie, Philip J. (2003). “A diverse dinosaur-bird footprint assemblage from the Lance Formation, Upper Cretaceous, eastern Wyoming; implications for ichnotaxonomy”. Ichnos, 11: 229-249.
  • Lockley, Martin; Gierlinski, Gerard; Adach, Lidia; Schumacher, Bruce; Cart, Ken (2018). “Newly Discovered Tetrapod Ichnotaxa from the Upper Cretaceous Blackhawk Formation, Utah”. In Spencer G. Lucas and Robert M. Sullivan, eds. New Mexico Museum of Natural History and Science. Fossil Record 6, Volume 2: Bulletin 79. Albuquerque: New Mexico Museum of Natural History and Science, 2018. Pages 469-480.
  • McCrea, R. T.; Buckley, L. G.; Plint, A. G.; Currie, Philip J.; Haggart, J. W.; Helm, C. W.; Pemberton, S. G. (2014). “A review of vertebrate track-bearing formations from the Mesozoic and earliest Cenozoic of western Canada with a description of a new theropod ichnospecies and reassignment of an avian ichnogenus”. In Lockley Martin G.; Lucas, Spencer G., eds. New Mexico Museum of Natural History & Science. Bulletin 62: Fossil Footprints of Western North America. Albuquerque: New Mexico Museum of Natural History & Science, 2014. Pages 5-94.
  • McCrea, R. T.; Tanke, D. H.; Buckley, L. G.; Lockley, Martin G.; Farlow, James O.; Xing, L.; Matthews, N. A.; Helm, C. W.; Pemberton, S. G.; Breithaupt, B. H. (2015). “Vertebrate ichnopathology: pathologies inferred from dinosaur tracks and trackways from the Mesozoic”. Ichnos, 22 (3–4): 235-260.
  • Zanno, Lindsay Elizabeth. A Taxonomic and Phylogenetic Reevaluation of Therizinosauria (Dinosauria: Theropoda): Implications for the Evolution of Maniraptora. PhD dissertation, submitted to the University of Utah. December 2008.

Websites:

Videos:

  • Utah’s Dino Graveyard. The Discovery Channel, 2005.
  • When Dinosaurs Roamed America. The Discovery Channel, 2001.

 

Hybodus, the iconic shark of the dinosaur age

Many people, usually un-informed talking heads that appear on populist nature documentaries who want to make claims that will grab your attention, will say that sharks have remained unchanged since the time of the dinosaurs. It’s wrong. The Mesozoic Era, the age of the dinosaurs, was a time of great transition for sharks. Sharks had existed on Earth for millions of years before the first dinosaurs appeared, ever since the Devonian Period when creatures like Cladoselache swam in the oceans around 370 million years ago. However, these were very primitive sharks that bore only a slight resemblance to most of the sharks that are found in the oceans today. The closest visual comparisons that we have for many prehistoric species are those that are found in very deep water, even though these sharks are still thoroughly modern in their genetics and evolutionary history. Sharks that are described as “modern” by biologists and paleontologists appeared towards the end of the Mesozoic Era during the late Cretaceous Period. Examples of prehistoric “modern sharks” are Cretoxyrhina and Squalicorax, both of which look like many shark species that are alive today.

During the Mesozoic Era, new shark forms emerged that could be described as transitional, a sort of in-between stage between the primitive sharks of the Paleozoic Era and the modern sharks of the very late Mesozoic. The most recognizable of these transitional species were a group of sharks called the “hybodonts”. Part of the reason why the hybodonts are regarded by many as THE shark group of the dinosaur age is because they lasted for such a long time. The hybodonts first emerged during the Carboniferous Period and they stuck around until the very end of the Cretaceous – that’s a LONG time! The hybodonts, therefore, existed throughout the entire duration of the Mesozoic Era. No wonder that they are considered the archetypal Mesozoic shark. Another reason for their status as the sharks that most people think of as dinosaur-age sharks is that they were widespread. Hybodont sharks existed all over the world during the Mesozoic, and they appeared to have existed in every aquatic niche: freshwater, brackish, and saltwater. Perhaps, like modern-day Bull Sharks, they had the ability to migrate in and out of water with different salinity levels without suffering adverse effects.

The most well-known of all of the hybodont sharks is its eponymous member Hybodus, a genus composed of several species which survived and thrived during the dinosaur age. It measured 6 feet long, and it occupied marine habitats around the world, although it is especially known from fossils found in Europe. Below is a drawing that I made of it based upon numerous fossils and scientific articles that I found. I noticed that the profile of the creature looked remarkably similar to a modern-day Blunt-Nosed Six-Gill Shark (Hexanchus griseus), with its rounded blunt nose and the characteristic humped back. Hybodus’ pectoral fins were surprisingly small and convex along the posterior edge, looking similar to the pectoral fins on numerous species of bottom-dwelling sharks. I get the impression that Hybodus was somewhat lethargic and spent much of its time cruising near the sea bottom, but that’s just my guess. The drawing was made with No. 2 pencil.

Keep your pencils sharp.

Habrosaurus, a Late Cretaceous siren amphibian from the Hell Creek Formation

The Hell Creek Formation of the north-central United States is famous for its dinosaur fossils, notably those of Tyrannosaurus, Triceratops, and others whose names are well-known to children and adults. However, this fossilized environment was home to more than just dinosaurs. The Hell Creek Formation was home to a wide range of fish, amphibians, reptiles, and mammals. One of the animals that called this landscape home during the late Cretaceous was Habrosaurus.

Despite its name, Habrosaurus was not a dinosaur, and it wasn’t even a reptile. It was, in fact, an amphibian, and a large one at that. Habrosaurus dilatus was a three-foot long siren, a type of salamander that bears more of a resemblance to an eel than the lizard-like forms that we associate salamanders with. Unlike most salamanders, sirens are fully-aquatic amphibians that retain gills throughout their whole lives, unlike other amphibian species that possess gills only in the early development stages of their lives. Sirens also possess small rudimentary lungs, and are able to breath air. There are four species of sirens that are alive today, and all of them are found within North America. Depending upon the species, they can have one to three gill slits on each side of the head. They have completely lost their hind limbs, and their front limbs have shrunk considerably, with three or four short stubby fingers on each hand. Sirens have tiny eyes and no eyelids, and possess a long tail reminiscent of an eel or a sea snake – ideal for swimming. Sirens prefer to live in slow-moving or static bodies of water with lots of underwater vegetation and muddy bottoms. They might occasionally come onto land during the night if the ground is wet or if it’s raining.

Habrosaurus is, to date, the oldest-known siren genus. So far, there are two species known: H. prodilatus, which was found in Alberta, Canada in rocks dating to the Campanian Phase (83-70 MYA) of the late Cretaceous, and H. dilatus, which is much more widespread in the western United States, being found in Montana and Wyoming (with more specimens being found in Wyoming) and dating to the Maastrichtian Stage (70-65 MYA) of the late Cretaceous, as well as being found in the early Paleocene Epoch of the Tertiary Period. This means that H. dilatus was one of several species to survive the K-T Extinction, if only for a short while. It may be possible that H. dilatus is simply the evolved form of H. prodilatus.

Habrosaurus dilatus was named by the eminent paleontologist Charles W. Gilmore in 1928. To my knowledge, six specimens have been found of this animal, and all of them have been found in stream channel deposits. The presence of this type of animal, as well as its impressive size of three feet in length, indicates the presence of large bodies of fresh water, such as slow-moving rivers or ponds. However, the possibility of a dry year was ever-present, and for a fully-aquatic or mostly-aquatic animal like Habrosaurus, that could spell doom. During dry periods or droughts, modern-day sirens are able to dig burrows into the mud and encase themselves in a cocoon, like a lungfish, and Habrosaurus might have adopted the same strategy.

Habrosaurus had rows of blunt teeth arranged in the roof of its upper jaw, which indicates that these jaws were designed for crushing rather than grabbing. Presumably, it fed upon tiny mollusks and arthropods, such as snails and shrimp. Modern-day sirens feed mainly upon worms, aquatic snails, shrimp, and occasionally algae. Like fish, sirens possess lateral lines to find prey by indicating differences in water pressure and underwater vibrations.

An appropriate modern-day analog for the three-foot long Habrosaurus dilatus is the Greater Siren (Siren lacertina), which also grows to three feet long and is the largest siren species in the world today.

Below is a simple drawing of a Habrosaurus that I made with a felt-tipped marker. This style is a considerable departure from my usual style of highly-detailed pencil drawings, but I wanted to do some artistic experimenting.

Dinosaur Day 2015 at the Garvies Point Museum

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Well, it was that time of year again! Every April or so, at around the time of Easter, the Garvies Point Museum and Preserve, located in Glen Cove, Nassau County, New York, holds it annual “Dinosaur Day”. This is one of the days that I really look foward to for a few reasons. First, I get to work at a place that I absolutely love and meet with some good friends. Secondly, I get to be out of NYC for a little while, which is something that I ALWAYS look foward to. Third, I get to talk about a subject that has fascinated me since my earliest days – paleontology.

Veronica, the museum’s de facto head of administration, did a wonderful job along with other members of the museum staff of setting up the classroom where the day’s major activities would be taking place. Recently, the museum’s library was substantially increased. The Sands Point Museum and Preserve had closed down its library a short while ago, and all of the books and papers were sent to the GPM. I should state, though, that almost all of these documents were originally part of the GPM collections anyway, and they just got them back, that’s all. However, Louis (one of the workers at the Garvies Point Museum, but works primarily at the Old Bethpage Village – another place that I really love) has been working hard to re-catalogue all of these books and papers back into the museum’s database.

The name of the event was somewhat misleading, as it concerned all prehistoric life, not just dinosaurs. We had exhibits on primitive mammal-like-reptiles, dinosaurs, and prehistoric mammals.

Here are some pictures of what the room looked like both during and after the hoards of kids showed up.

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Most of the really young children gravitated immediately towards the dino toy area and the fossil digsite. The older children and a lot of the adults were interested in the information that I and others were giving. They were especially interested in Dimetrodon, the famous sail-backed pelycosaur from the early Permian Period. I don’t think that I have ever had to say the name”Dimetrodon” so many times within the course of a single day! It seemed to be the only thing that many of them wanted to talk about!

Some of the major topics of interest on this day were: the Permian Mass Extinction, which occured about 251 million years ago, when an estimate 95% of all life was wiped out; of course, T. rex was a favorite; as too was Allosaurus, who competed with its larger relative for attention from the crowds. This was helped in no small part to the fact that we had a lot of Allosaurus “stuff” arrayed for them: a picture of the skull, a hand model, bone casts, a model, and my drawing which you might recognize from an earlier post on this blog.

Finally, here’s a picture of me, “the Dinosaur Man” as several members of the museum staff call me, dressed up as an amateur paleontologist. In addition to my olive drab Garvies Point Museum shirt, I also wore a khaki utility vest, because apparently ALL paleontologists wear khaki utility vests! I thought that wearing it would help to enhance my ethos with the audience, and by my reckoning, it worked.

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Lonchidion, a prehistoric shark

Lonchidion

This is a drawing of Lonchidion, a hybodont shark from the Mesozoic Era. There were at least eleven different species, one of which was found in the Hell Creek Formation. I won’t get into all of the particulars regarding this genus or the Hell Creek species in particular (L. selachos). Their size depended upon the species, some being very small. Lonchidion selachos may have been three feet long, judging by the size of its dorsal spines. The drawing is based upon the preserved remains of other hybodont sharks, because specimens from the Hell Creek Formation consist mostly of teeth, well-preserved specimens of any Lonchidion species are very rare, and as far as I am aware, they looked more or less like other well-known hybodonts.

Hybodont sharks are identified by their large dorsal fin spines as well as the four large spines atop their heads, which are really overly-enlarged denticle scales found all over the rest of the body. Hybodonts first appeared during the Carboniferous Period, but it was during the Jurassic that they came into their own. However, by the Cretaceous Period, they were being replaced by so-called “modern” sharks very similar to the ones we see today. Lonchidion was one of the last surviving members of its kind before the whole hybodont group (the few species that remained, anyway) was completely wiped out at the end of the Mesozoic Era 65 million years ago.

News: “Nanook of the North”? New tyrannosaur species from Alaska

Paleontologists have recently announced the discovery and naming of a new tyrannosaur species from Alaska. They have called it Nanuqsaurus hoglundi.

The discovery was made by a team of paleontologists working for the Perot Museum of Nature and Science, located in Dallas, Texas; the team was led by Prof. Anthony R. Fiorillo. The fossils were found at the Prince Creek Formation in Alaska in 2006 when the team was hunting for ceratopsians (that’s “horned-faced” dinos, like Triceratops and Styracosaurus). They consist of the front portion of the lower jaw (the bone is called the “dentary” because that’s the bone in the lower jaw that has the teeth in it) and two small pieces of the upper jaw. The pieces were collected, and then gathered dust for a while until Prof. Fiorillo and his associate Dr. Ronald Tykoski re-examined them.

Admitedly, it’s not that much to go on, but apparently, it was enough to create not only a new species, but a new genus. That doesn’t surprise me at all, as paleontologists are well known to be afflicted with what I call “neogenitis” – “the new genus disease”. They just can’t resist making up new names for things. The name Nanuqsaurus derives from the Inupiak word nanuq, meaning “polar bear”, and the ancient Greek word sauros, “lizard”. The species name is in honor of the philanthropist Forrest Hoglund.

The fossils date to 70 million years ago. It appears to be closely related to both Tyrannosaurus rex and a close relative called Tarbosaurus bataar which lived in Mongolia (some paleontologists consider Tarbosaurus bataar merely to be an Asian species of Tyrannosaurus – personally, I don’t buy it for a few reasons, but I won’t get into them here). Based upon the size of the remains, limited though they may be, Nanuqsaurus may have been only half the size of T. rex.

Professor Fiorillo suspects that the animal’s small size might be a reference to a limited food supply up in the Great un-White North of the late Cretaceous. Although only three small pieces were recovered, it is strongly plausible that a northern tyrannosaur like Nanuqsaurus would be covered in an insulating layer of feathery fuzz.