One of the world’s most unusual dinosaurs is even stranger than first thought.  Newly published research in the journal “Nature” confirms that the Moroccan armoured dinosaur Spicomellus afer is definitely an ankylosaur.  In addition, to the extremely spiky appearance, it probably had a tail weapon.  The evolution of a tail weapon predates this feature in any other known ankylosaur by more than thirty million years.  Furthermore, Spicomellus had a unique bony collar ringed with metre-long spikes sticking out from either side of its neck.

It has been nicknamed the “punk rock dinosaur”.

A Spicomellus life reconstruction in anterior view. Picture credit: Matthew Dempsey.

Picture credit: Matthew Dempsey

This dinosaur was formally named and described in 2021 (Maidment et al).  The initial description was made based on a single rib bone. The rib had spikes fused to it, a unique feature, not seen in any other animal.  However, the “T-shaped” cross section of the rib permitted the scientists to confidently assign this fossil to an ankylosaur.  Named Spicomellus afer, it represents Africa’s first known ankylosaur and the earliest representative of this iconic dinosaur clade (Ankylosauria).

To read Everything Dinosaur’s blog post from 2021 announcing the discovery of Africa’s first ankylosaur: The Earliest Ankylosaur and Africa’s First – Spicomellus.

The Remarkable Early Ankylosaur Spicomellus afer

The fossils are more than 165 million years old. This armoured dinosaur lived during the Middle Jurassic, near what is now the Moroccan town of Boulemane.  Further fossil discoveries have enabled the research team to learn more about this remarkable armoured dinosaur.  For example, they now know that Spicomellus had bony spikes fused onto and projecting from all of its ribs, a feature not seen in any other vertebrate species living or extinct. It had long spikes, measuring eighty-seven centimetres in length, which the researchers believe would have been even longer in real life.  These spikes emerged from a bony collar that sat around the reptile’s neck.

A fossil rib showing the spikes fused to it, a unique feature not seen in any other animal. Picture credit: The Trustees of the Natural History Museum, London.

Picture credit: The Trustees of the Natural History Museum, London

Professor Susannah Maidment of Natural History Museum, London, and the University of Birmingham, who co-led the team of researchers commented:

“To find such elaborate armour in an early ankylosaur changes our understanding of how these dinosaurs evolved. It shows just how significant Africa’s dinosaurs are, and how important it is to improve our understanding of them.”

Elaborate Dermal Armour

Ankylosaurs are best known from Late Cretaceous Northern Hemisphere ecosystems.  For instance, Ankylosaurus and Euoplocephalus are known from Upper Cretaceous rocks in the northern United States and Canada.  Ziapelta is known from fossils found in New Mexico, whereas Saichania, Pinacosaurus and Tarchia are known from Upper Cretaceous rocks in Asia.

“Sede” the Ankylosaurus dinosaur model.  The authors of the Spicomellus study postulate that with the emergence of larger predators this could have resulted in ankylosaur armour becoming simpler and more defensive.  Picture credit: Everything Dinosaur.

Picture credit: Everything Dinosaur

The picture (above) shows a model of the Late Cretaceous ankylosaur Ankylosaurus magniventris.  The figure is from the Chinese manufacturer PNSO.

To view the range of PNSO models and figures in stock: PNSO Age of Dinosaurs Models.

The researchers postulate that the unique, elaborate spines and spikes of Spicomellus may have functioned for display as well as defence.  Later ankylosaurs had simpler armour with less extravagant osteoderms.  This might indicate a shift towards a primarily defensive function, perhaps in response to increased predation pressures or a switch to combative courtship displays.

Professor Maidment added:

“Spicomellus had a diversity of plates and spikes extending from all over its body, including metre-long neck spikes, huge upwards-projecting spikes over the hips, and a whole range of long, blade-like spikes, pieces of armour made up of two long spikes, and plates down the shoulder. We’ve never seen anything like this in any animal before. It’s particularly strange as this is the oldest known ankylosaur, so we might expect that a later species might have inherited similar features, but they haven’t.”

Professor Susannah Maidment of the Natural History Museum holding a rib with fused spikes. Picture credit: The Trustees of the Natural History Museum, London.

Picture credit: The Trustees of the Natural History Museum, London

Was There a Large Theropod in the Ecosystem?

There is another potential explanation for the remarkable armour associated with Spicomellus afer.  Could it have shared its environment with a large predator such as a theropod dinosaur?  There is certainly evidence to suggest that by the Middle Jurassic formidable tetanuran theropods were present in many ecosystems.

Co-author of the study, Professor Richard Butler (University of Birmingham) stated:

“Seeing and studying the Spicomellus fossils for the first time was spine-tingling. We just couldn’t believe how weird it was and how unlike any other dinosaur, or indeed any other animal we know of alive or extinct. It turns much of what we thought we knew about ankylosaurs and their evolution on its head and demonstrates just how much there still is to learn about dinosaurs”.

Professor Susannah Maidment of the London Natural History Museum and Professor Richard Butler (University of Birmingham) examine the fossil remains along with fellow researchers. Picture credit: The Trustees of the Natural History Museum, London.

Picture credit: The Trustees of the Natural History Museum, London

Did Spicomellus afer Have a Tail Club?

One feature of early ankylosaurs that may have survived, however, is their tail weaponry. While the end of Spicomellus’ tail has not be found, the caudal vertebrae that do survive suggest that it had a club or a similar tail weapon. Some of these tail vertebrae are fused together.  They form a structure referred to as a “handle”.  This feature has only been found in ankylosaurs that possessed a tail club.  If Spicomellus did have a tail club, it overturns current understanding regarding tail club evolution in the Ankylosauria.  These structures were previously thought to have first evolved in the Early Cretaceous.

The authors believe that the combination of a tail weapon and an armoured shield that protected the hips suggest that many of the ankylosaurs’ key adaptations already existed by the time of Spicomellus.

Spicomellus fossil material. This dinosaur was originally described in 2021, however, more fossils were excavated in 2023 providing the research team with further information about the bizarre anatomy of Spicomellus. Picture credit: The Trustees of the Natural History Museum, London.

Picture credit: The Trustees of the Natural History Museum, London

Improving Our Understanding of the Geographic Distribution of Armoured Dinosaurs

Finding more fossils of Spicomellus confirms its ankylosaurian affinities.  In addition, it helps to deepen our understanding of the geographic distribution of armoured dinosaurs. It also helps to spark public imagination in the Dinosauria as we learn more about the baffling characteristics of species like Spicomellus.

Professor Driss Ouarhache, lead of the Moroccan team from the Université Sidi Mohamed Ben Abdellah who co-developed the research, commented:

“This study is helping to drive forward Moroccan science. We’ve never seen dinosaurs like this before, and there’s still a lot more this region has to offer.”

The Spicomellus afer fossils that form the basis of this study were cleaned and prepared at the Department of Geology of the Dhar El Mahraz Faculty of Sciences in Fez, Morocco, using scientific equipment provided by the University of Birmingham’s Research England International Strategy and Partnership Fund. The fossils are now catalogued and stored on this site.  Perhaps, they will be put on display so that the public will have the opportunity to view these amazing fossils.

Everything Dinosaur acknowledges the assistance of a media release from the University of Birmingham in the compilation of this article.

In addition, Everything Dinosaur acknowledges the assistance of the London Natural History Museum for the use of their images.

The scientific paper: “Extreme armour in the world’s oldest ankylosaur” by Susannah C. R. Maidment, Driss Ouarhache, Kawtar Ech-charay, Ahmed Oussou, Khadija Boumir, Abdessalam El Khanchoufi, Alison Park, Luke E. Meade, D. Cary Woodruff, Simon Wills, Mike Smith, Paul M. Barrett and Richard J. Butler published in the journal Nature.

The award-winning Everything Dinosaur website: Everything Dinosaur.

The Queensland-based Age of Dinosaurs Natural History Museum has announced a remarkable Australian ichthyosaur discovery. The specimen, probably representing Platypterygius australis is perhaps the most complete ichthyosaur ever found “Down Under”.  It was unearthed some sixty miles south of McKinlay (Queensland).  Preparation will soon commence at the Australian Age of Dinosaurs Museum of Natural History.  The fossil find highlights the rich palaeontological heritage of this part of western Queensland.

An aerial view of the ichthyosaur quarry. Picture credit: Australian Age of Dinosaurs Museum.

Picture credit: Australian Age of Dinosaurs Museum of Natural History

A Remarkable Australian Ichthyosaur Discovery

The skeleton measures an impressive 7.1 metres in length.  It was discovered on Toolebuc Station by neighbouring property owner and fossil enthusiast Cassandra Prince two years ago.  The specimen was carefully excavated the following year by Cassandra and her family. Remarkably well-preserved, the skeleton includes a complete vertebral column, intact left flipper, partial right flipper, rare hind flippers, partial caudal vertebrae and a nearly complete skull and torso.  Palaeontologists have described this find as one of the most scientifically valuable marine reptile fossils from Australia.

Cassandra Prince with her ichthyosaur discovery. Picture credit: Australian Age of Dinosaurs Museum.

Picture credit: Australian Age of Dinosaurs Museum of Natural History

Platypterygius australis

The Platypterygius genus is a geographically and temporally widespread genus.  Numerous species have been named.  Although, their taxonomic affinity is controversial all the specimens assigned to this genus are robust, macropredators with robust teeth.  It is likely that Platypterygius australis was a formidable predator in the inland sea (Eromanga Sea) that covered much of Australian in the late Early Cretaceous.

Museum Founders David and Judy Elliott assisted Cassandra and her family with collecting the specimen, which was subsequently transported to the Museum so that its preparation can start.  The fossil specimen has been generously donated to the Museum by the Toolebuc Station owners.  Once cleaned and prepared, the ichthyosaur specimen will be mounted in a special exhibition at the Museum.

David Elliott stated:

“This find is a huge win for science and public exhibitions in Australia. Its discovery is testament to the dedication and expertise of Cassandra and her fossil-hunting family and the unique geological heritage of the region. We look forward to sharing this incredible piece of Australia’s prehistory with visitors for generations to come.”

The Toolebuc fossil ichthyosaur specimen uncovered. This 7.1 metre specimen represents a remarkable Australian ichthyosaur discovery. Picture credit: Australian Age of Dinosaurs Museum.

Picture credit: Australian Age of Dinosaurs Museum of Natural History

Everything Dinosaur acknowledges the assistance of a media release from the Australian Age of Dinosaurs Museum in the compilation of this article.

The award-winning Everything Dinosaur website: Prehistoric Animal Models and Toys.

Researchers have described a new dinosaur species from the Wessex Formation of the Isle of Wight.  The dinosaur, an iguanodontian, has been named Istiorachis macarthurae.  The extended neural spines associated with the dorsal and caudal vertebrae suggest that it possessed a sail structure.

Possible explanations for neural spine elongation in the Ankylopollexia include biomechanical advantage, perhaps related to increasing body size and a move towards quadrupedalism. In addition, such structures could have evolved as aids to visual signalling or to deter rivals. Perhaps these changes in body shape were driven by sexual selection, species recognition or both

Hyperelongation of neural spines is known in a number of dinosaur taxa. Most recorded incidences occur during the Barremian and early Aptian faunal stages of the Early Cretaceous. The evolution of elongated neural spines probably took place due to a variety of evolutionary pressures. The drivers for this body shape probably differed in different taxa. Furthermore, it is likely that no single explanation fully supports the variation seen throughout the Cretaceous.

A life reconstruction of Istiorachis macarthurae. Picture credit: James Brown (University of Portsmouth).

Picture credit: James Brown (University of Portsmouth)

Istiorachis macarthurae

This new iguanodontian dinosaur was identified by Jeremy Lockwood, a retired GP, as part of his PhD studies at the University of Portsmouth and the Natural History Museum, London.  Detailed analysis of fossil bones held in the collection of the Dinosaur Isle museum, Isle of Wight led to the identification of several autapomorphies which resulted in the establishment of this new taxon.

The genus name is derived from the Ancient Greek words ἱστίον (istion), meaning a sail, and ῥάχις (rachis), the spine or backbone. It refers to the probable sail-back appearance of the dinosaur.

Pronounced Is-tree-oh-rak-is mack-ar-four-eye, the species name honours the Isle of Wight resident Dame Ellen MacArthur. Dame Ellen MacArthur is a famous sailor.  Therefore, it seemed appropriate to honour her by naming a possible sail-backed dinosaur after her. In 2005, she set a world record for the fastest solo non-stop voyage around the world on her first attempt and Dame Ellen MacArthur has also founded the Ellen MacArthur Cancer Trust for young people on the Isle of Wight.

Dr Jeremy Lockwood with the spinal column of Istiorachis macarthurae with some of the pelvic elements (pubis and head of the ischium). Note the elongated neural spines. Picture credit: University of Portsmouth.

Picture credit: University of Portsmouth

Smaller Iguanodontians Including Istiorachis macarthurae

Dr Lockwood has played a significant role in helping palaeontologists to better understand the smaller iguanodontians from the Isle of Wight.  This is the third iguanodontian named from fossils found on the island in recent years.  For example, last year (2024), Comptonatus chasei was described.  Moreover, in 2021, an iguanodontian with an unusual bulbous snout was described (Brighstoneus simmondsi).

To read Everything Dinosaur’s blog post about the discovery of Comptonatus chasei: A New Dinosaur From the Isle of Wight.

To learn more about B. simmondsi: A New Iguanodontid from the Isle of Wight.

A Highly Diverse Early Cretaceous Ecosystem

The fossil material is estimated to be around 125 million years old.  It came from Wessex Formation exposures from the southwestern part of the island.  This discovery further demonstrates the remarkable dinosaur diversity during the Early Cretaceous.  In addition, it helps to cement the Isle of Wight as a globally significant location for dinosaur fossils.

Commenting on how the new taxon was established, Dr Lockwood explained:

“While the skeleton wasn’t as complete as some of the others that have been found, no one had really taken a close look at these bones before. It was thought to be just another specimen of one of the existing species, but this one had particularly long neural spines, which was very unusual.”

The findings have been published this week in the academic journal “Papers in Palaeontology”.

Dr Lockwood holding the single cervical vertebra (neck bone) known from Istiorachis macarthurae. Picture credit: University of Portsmouth.

Picture credit: University of Portsmouth

Dr Lockwood added:

“Evolution sometimes seems to favour the extravagant over the practical. While the exact purpose of such features has long been debated – with theories ranging from body heat regulation to fat storage – researchers believe that the most likely explanation in this case is visual signalling, possibly as part of a sexual display and this usually is because of sexual selection. In modern reptiles, sail structures often show up more prominently in males, suggesting that these attributes evolved to impress mates or intimidate rivals. We think Istiorachis may have been doing much the same.”

A Detailed Analysis of Neural Spines in Iguanodontids and Their Relatives

A large database was constructed consisting of data on the neural spines of iguanodontids and their close relatives.  The database was created using observations, photographs, scientific illustrations and reconstructions of vertebrae.  The data enabled the researchers to trace the evolutionary history of elongated neural spines within the Iguanodontia.  This analysis permitted the team to identify broad trends in the evolution of sail-like structures.

Dr Lockwood stated:

“These methods let us move beyond simply describing the fossil and actually test hypotheses about its function. We showed that Istiorachis’s spines weren’t just tall – they were more exaggerated than is usual in Iguanodon-like dinosaurs, which is exactly the kind of trait you’d expect to evolve through sexual selection.”

Co-author of the study, Professor Susannah Maidment (London Natural History Museum), added:

“Jeremy’s careful study of fossils that have been in museum collections for several years has brought to life the iguanodontian dinosaurs of the Isle of Wight. His work highlights the importance of collections like those at Dinosaur Isle, where fossil specimens are preserved in perpetuity and can be studied and revised in the light of new data and new ideas about evolution. Over the past five years, Jeremy has single-handedly quadrupled the known diversity of the smaller iguanodontians on the Isle of Wight, and Istiorachis demonstrates we still have much to learn about Early Cretaceous ecosystems in the UK.”

A silhouette of Istiorachis macarthurae showing known fossil material and providing a size estimate. Note scale bar = 50 cm. Picture credit: James Brown (University of Portsmouth) with additional annotation by Everything Dinosaur.

Picture credit: James Brown (University of Portsmouth) with additional annotation by Everything Dinosaur

Highlighting a Broader Evolutionary Trend

Importantly, Istiorachis macarthurae appears to highlight a broader evolutionary trend.  The database suggests that elongation of neural spines in iguanodontians began in the Late Jurassic.  It becomes an increasingly common feature during the Early Cretaceous. However, true hyper-elongation, where neural spines reach heights in excess of four times the height of the vertebral body remain rare.

Similar elongated spines are seen in living reptiles.  For example, many species of extant lizards sport elaborate crests and sail-like structures.  These play a role in visual communication as well as signalling the health and vitality of the animal.

Everything Dinosaur acknowledges the assistance of a media release from the University of Portsmouth in the compilation of this article.

The scientific paper: “The origins of neural spine elongation in iguanodontian dinosaurs and the osteology of a new sail-back styracosternan (Dinosauria, Ornithischia) from the Lower Cretaceous Wealden Group of England” by Jeremy A. F. Lockwood, David M. Martill, Susannah C. R. Maidment published in Papers in Palaeontology.

The award-winning Everything Dinosaur website: Prehistoric Animal Models and Toys.

Locomotion in the Sauropoda is hotly debated by palaeontologists.  Some of these leviathans grew to enormous sizes. Once thought to have been semi-aquatic, scientists have a better understanding of their anatomy, and most academics concur that these reptiles were well-adapted to a terrestrial existence. How these animals moved their enormous bulk has been the subject of numerous scientific papers.  In a new study, researchers have examined the caudal vertebrae of the giant Giraffatitan brancai to gain a better understanding of tail mobility.  This research, published in the journal “Royal Society Open Science” provides fresh insights into the biomechanical properties of one of the largest dinosaurs known to science.

The study analysed eighteen preserved tail vertebrae from a nearly complete tail of Giraffatitan brancai.  Excavated from Upper Jurassic deposits in Tanzania, these fossils are in the collection at the Museum für Naturkunde Berlin.

A model of Giraffatitan brancai. The W-Dragon Giraffatitan Compared to a Papo standing T. rex dinosaur model. Picture credit: Everything Dinosaur.

Picture credit: Everything Dinosaur

The tails of archosaurs are important.  They have a biomechanical function and assist with movement.  In addition, they function as behavioural tools helping animals to communicate and interact with their environment. Until recently, tails have been neglected in biomechanical analyses and were considered as a stiff (sometimes independent) unit. However, the tail’s role in movement is now increasingly being appreciated.

Testing the Movement of Individual Giraffatitan brancai Caudal Vertebrae

New kinematic programmes were used to test the movement of the individual caudal vertebra in relation to each other. The scientists wanted to explain the impact of forces on the muscles.  Furthermore, they wanted to pinpoint areas of muscle attachment on the tail bones.  From this, they set out to determine the mobility and movement of the dinosaur.

Corresponding author, Dr. Verónica Díez Díaz (Museum für Naturkunde Berlin), explained:

“Our analyses show that the tail of Giraffatitan was much more mobile and functionally versatile than previously assumed.”

A detailed analysis of the so-called haemal arches was undertaken.  These haemal arches are bony structures on the underside of the tail vertebrae. They were often overlooked in earlier studies.  In this new study, the researchers conclude that these elements had a significant influence on the mobility of the tail.

Diagram of specimen number MB.R.2921 caudal series of G. brancai, left, with (green) intervertebral discs between vertebrae. Colour coded computer model (right) testing the flexibility of the caudal vertebrae. Picture credit: Díez Díaz et al.

Picture credit: Díez Díaz et al

Employing Sophisticated Computer Models

The use of sophisticated computer models confirm that the Giraffatitan exhibit at the Berlin Natural History Museum is accurate.  Sauropods did not drag their tails behind them like crocodilians.  Instead, the carried their tails raised off the ground. Furthermore, Giraffatitan could move its tail flexibly in several directions.

This study provides new information on the posture, movement and possibly the social interactions of these massive dinosaurs. The study not only offers new perspectives on the anatomy of macronarian sauropods, it also provides valuable impetus for future reconstructions and palaeobiological interpretations.

Everything Dinosaur acknowledges the assistance of a media release from the Museum für Naturkunde Berlin in the compilation of this article.

The scientific paper: “Centres of rotation and osteological constraints on caudal ranges of motion in the sauropod dinosaur Giraffatitan brancai” by Verónica Díez Díaz, Pasha A. van Bijlert, William Irvin Sellers, Mathew J. Wedel and Daniela Schwarz published in Royal Society Open Science.

The award-winning Everything Dinosaur website: Dinosaur Models and Figures.

A new species of Early Jurassic plesiosaur has been named based on a detailed examination of an almost complete fossil specimen discovered nearly fifty years ago.  The articulated specimen represents a skeletally immature individual, and it has been named Plesionectes longicollum.  Writing in the open-access journal PeerJ, the researchers identified several unique anatomical features of the fossil that permitted the erection of a new taxon. For instance, this plesiosaur had an exceptionally long neck.  At least forty-three cervical vertebrae have been identified.  The count hampered by the poor state of preservation of the skull.

The fossil material had been studied previously. For example, evidence of soft tissue preservation had been identified on the neck, tail and rear limbs. In addition, V-shaped neurocentral sutures have been identified.  Furthermore, this animal had between twenty and twenty-one dorsal vertebrae, one of the highest counts known among Lower Jurassic plesiosaurs.

Moreover, the fossil was excavated from a different horizon from the famous oil shales exposed near to Holzmaden. It is much older than other plesiosaurs known from the Posidonia Shale (Posidonienschiefer Formation). The remains of this marine reptile are thought to be around 183 million years old.

Plesionectes longicollum life reconstruction. Picture credit: Peter Nickolaus.

Picture credit: Peter Nickolaus

Plesionectes longicollum

Researcher Sven Sachs (Naturkunde-Museum Bielefeld, Germany) in collaboration with his colleague Daniel Madzia from the Polish Academy of Sciences, conducted the first detailed osteological examination of the fossil material.  This fossil is surprisingly large compared to other plesiosaurs known from the Posidonia Shale. It measures approximately 2.95 metres long, but when the head is included, this marine reptile would have measured around 3.2 metres in length.

The genus name derives from the Greek for “close” or “near”, referring to its plesiosaur affinities, and the Greek word for “swimmer” a common suffix in plesiosaur taxon nomenclature. The species name derives from the Latin and means “long neck”.

To read a recent blog post about examination of soft tissue preservation in a plesiosaur from the Posidonia Shale: Lower Jurassic Plesiosaur Soft Tissue is Examined.

The Plesionectes longicollum holotype (SMNS 51945). Patches of soft tissue have been preserved around the neck, tail, and hindlimb. This marine reptile is thought to have measured around 3.2 metres in length. Picture credit: Sven Sachs.

Picture credit: Sven Sachs

Discovered in 1978

The fossil material was found in 1978. It was discovered by Gotthilf Fischer in his own quarry in Holzmaden. The specimen was prepared by its discoverer and in 1979 it was acquired by the Staatliches Museum für Naturkunde (Stuttgart).

In personal correspondence with Everything Dinosaur, researcher Sven Sachs outlined the direction of his future studies. He intends to commence a reassessment of the famous genus Plesiosaurus.

Sven commented:

“It will surely take a few years to visit all the collections where Plesiosaurus material is housed, to analyse the data, and to write it all up, but this will be a fun project as it is such an iconic taxon.”

A close view of the posterior cervical vertebrae of Plesionectes longicollum. This early-diverging plesiosauroid from the Lower Jurassic Posidonia Shale of Holzmaden, Germany had at least 43 neck bones. Picture credit: Sven Sachs.

Picture credit: Sven Sachs

In summary, this newly described species helps to define the remarkable diversity of Lower Jurassic plesiosauroids known from Germany.  In addition, it invites renewed examination of specimens in museum collections.

Everything Dinosaur acknowledges the assistance of one of the study’s authors in the compilation of this article.

The scientific paper: “An unusual early-diverging plesiosauroid from the Lower Jurassic Posidonia Shale of Holzmaden, Germany” by Sven Sachs and Daniel Madzia published in PeerJ.

A team of international researchers have named a new species of Permian archosauromorph based on a single neck bone.  The animal has been named Manistropheus kulicki.  It provides new insights into the early evolution of archosauromorphs, a clade that includes the crocodiles, birds, pterosaurs and dinosaurs.  The cervical vertebra was discovered last century, but it has only just been scientifically described.  It was found at the famous Korbacher Spalte site in central Germany.  A fissure preserves the fragmentary and isolated remains of many Late Permian vertebrates.  The sediments are believed to around 255 million years old.

The Korbacher Spalte locality is important because it preserves evidence of tetrapods prior to the mass extinction event at the Permian-Triassic boundary.

The holotype of Manistropheus kulicki (SMNK-PAL 76022) shown in left lateral view. Picture credit: Carola Radke.

Picture credit: Carola Radke

Korbacher Spalte

The Korbacher locality is particularly well known for its many finds of the early mammal ancestor, a cynodont called Procynosuchus.  This synapsis is also jokingly referred to as the “Korbach dachshund” because of its appearance. However, scientists have now described a previously unknown species of archosauromorph reptile based on a single, well-preserved cervical vertebra. Distinctive characteristics of the fossil bone enabled the team to erect a new genus and species – Manistropheus kulicki.

Manistropheus kulicki Cervical Vertebra

The vertebra is characterised by an elongated, diamond-shaped centre and a crescent-shaped indentation on the side of the front edge of the vertebra. This gives the new genus its name – from the Old Norse Máni, the personification of the moon in Germanic mythology, and the Greek “stropheus,” meaning vertebra. Overall, the specimen shows similarities to early archosauromorphs but also shows features that are absent in other reptiles of that time. A comprehensive study of the phylogenetic relationships suggests that M. kulicki stands at the base of this important reptile lineage.  It is thought to be a basal archosauromorph.

The study also used an analysis of morphological diversity to investigate how cervical vertebrae have changed over the course of Earth’s history. The results suggest that archosauromorphs were already morphologically diverse before their extinction and that their cervical anatomy diversified rapidly in the Early Triassic.  Cervical vertebrae anatomy changed faster than other parts of the skeleton.

Lead author of the study, Dr Martín Ezcurra (CONICET) stated:

“This discovery is particularly significant because Permian archosauromorphs are extremely rare, with only five fossil species from this period known to date. Thanks to Manistropheus kulicki, we can see how diverse this group already was before the mass extinction.”

Co-author Professor Hans-Dieter Sues (Smithsonian Institute), added:

“This fossil not only proves the existence of a new species, but also supports the assumption that there was already a previously hidden diversity of archosauromorphs in the Permian period.”

Diverse Archosauromorphs Present in Equatorial Regions During the Late Permian

Professor Jörg Fröbisch (Museum für Naturkunde Berlin), another co-author of the study, highlighted the significance of the Korbach Spalte site.

He commented:

“The Korbach fissure site is proving to be a key location for better understanding life on land in the tropical regions of the supercontinent Pangaea shortly before the largest mass extinction in Earth’s history. “

The naming of this basal archosauromorph from Germany highlights the importance of continuing to explore lesser-known fossil sites.  It is especially important to explore fossil sites that provide insights into ancient ecosystems threatened with extinction.

Everything Dinosaur acknowledges the assistance of a media release from the Museum für Naturkunde Berlin in the compilation of this article.

The scientific paper: “A new late Permian archosauromorph reptile from Germany enhances our understanding of the early diversity of the clade” by Martín D. Ezcurra, Hans-Dieter Sues and Jörg Fröbisch published in the Journal of Systematic Palaeontology.

The Everything Dinosaur website: Prehistoric Animal Models.

Stunning fossils of Cambrian nectocaridids solves a half-a-billion-year-old evolutionary puzzle.  Fossil material excavated from the Sirius Passet site in North Greenland has enabled palaeontologists to identify these ancient marine invertebrates as early descendants of chaetognaths (arrow worms).  It had been proposed that these animals were a type of cephalopod.  Cephalopods are marine molluscs with tentacles and a prominent head.  Living cephalopods include cuttlefish, octopi and squid.  The extinct ammonites and belemnites were cephalopods.

This significant discovery rewrites part of the story of the Cambrian Explosion. It shows arrow worms descended from complex predators, not simple marine organisms.

Life reconstruction of Nektognathus, swimming in the Cambrian Sea. It is depicted as a nektonic predator. Picture credit: Bob Nicholls.

Picture credit: Bob Nicholls

The illustration of Cambrian life (above) was created by acclaimed palaeo-reconstruction artist Bob Nicholls.  Bob has recently been collaborating with Everything Dinosaur in the production of museum quality prehistoric animal models – Everything Dinosaur Evolution.

To learn more about these remarkable, scientifically accurate figures: Everything Dinosaur Evolution Figures.

Studying Cambrian Nectocaridids

The research team included palaeontologists from the University of Bristol, the University of Copenhagen, and the Korean Polar Research Institute. Their work builds on nine years of expeditions to Sirius Passet, a site famous for exceptional soft-tissue fossil preservation from 518 million years ago (Stage 3 of the Cambrian).

Co-author of the study, Dr Jakob Vinther, Associate Professor in Macroevolution at the University of Bristol commented:

“Sirius Passet is a treasure trove of fossils from the Cambrian Explosion. We not only find delicate soft-bodied fossils but also their digestive systems, musculature and sometimes even their nervous system.”

Research first published in 2009 examining fossils from the famous Burgess Shale deposits of British Columbia (Canada), linked nectocaridids to cephalopods.  However, some invertebrate palaeontologists remained sceptical.

Dr Vinther explained:

“Around 15 years ago a research paper, based on fossils from the famous Burgess Shale, claimed nectocaridids were cephalopods. It never really made sense to me, as the hypothesis would upend everything we otherwise know about cephalopods and their anatomy didn’t closely match cephalopods when you looked carefully.”

Field team members became excited as they found twenty-five new specimens of Cambrian nectocaridids in the Sirius Passet deposits. These are the first nectocaridids unearthed from the Sirius Passet locality. Evidence of nervous systems were preserved in some specimens.

A fossil specimen of Nektognathus discovered at Sirius Passet (northern Greenland). Picture credit: University of Bristol.

Picture credit: University of Bristol

Identifying the Ventral Ganglion

The team discovered nectocaridids with parts of their nervous system preserved as paired mineralised structures.  This new evidence helped to confirm the phylogeny of these ancient animals. Arrow worm fossils with preserved ventral ganglia – a unique nerve structure seen in modern chaetognaths (arrow worms) were also found.

Co-author of the paper, Dr Tae-Yoon Park from the Korean Polar Institute stated:

“These fossils all preserve a unique feature, distinct for arrow worms, called the ventral ganglion.”

The discovery of this structure confirms the evolutionary link between nectocaridids and arrow worms.

The holotype specimen of Nektognathus from Sirius Passet. Picture credit: Tae-Yoon Park.

Picture credit: Tae-Yoon Park

The authors of the study have erected a new nectocaridid taxon Nektognathus evasmithae. The species name honours Professor Eva Smith, the first female professor of law in Denmark and renowned human rights advocate. Dr Vinther commented that N. evasmithae was a smart and stealthy fighter, just like the Danish advocate.

Dr Park added:

“We now had a smoking gun to resolve the nectocaridid controversy. Nectocaridids share a number of features with some of the other fossils that also belong to the arrow worm stem lineage. Many of these features are superficially squid-like and reflect simple adaptations to an active swimming mode of life in invertebrates, just like whales and ancient marine reptiles end up looking like fish when they evolve such a mode of life.”

The holotype specimen of Nektognathus from Sirius Passet imaged with Electron Probe microanalysis for its carbon content. Picture credit: Mirinae Lee and Chankun Park.

Picture credit: Mirinae Lee and Chankun Park

Cambrian Nectocaridids – Dynamic, Nektonic, Agile Predators

This surprising discovery means the rather simple marine arrow worms had ancestors with much more complex anatomies and a predatory role higher up in the food chain. Cambrian nectocaridids had camera-type eyes, long antennae, and streamlined bodies for active hunting.  In contrast, extant arrow worms have eyes that are sensitive to light but are not thought to have effective vision.  The researchers conclude that the ancestors of today’s arrow worms were nektonic predators.  They filled an ecological niche similar to living squid.  Although, squids evolved much later.  Preserved gut contents indicate that Nektognathus evasmithae fed on the swimming arthropod Isoxys.

Dr Vinther commented:

“We can therefore show how arrow worms used to occupy a role much higher in the food chain. Our fossils can be much bigger than a typical living arrow worm and combined with their swimming apparatus, eyes and long antennae, they must have been formidable and stealthy predators.”

A photograph of a modern chaetognath (arrow worm) from Antarctica. Picture credit: Tae-Yoon Park.

Picture credit: Tae-Yoon Park

This study gives us a new view of Cambrian nectocaridids. Far from being early squids, they were fierce predators with an advanced anatomy.  This study helps palaeontologists to better understand Cambrian marine ecosystems.

Everything Dinosaur acknowledges the assistance of a media release from the University of Bristol in the compilation of this article.

The scientific paper: “A fossilised ventral ganglion reveals a chaetognath affinity for Cambrian nectocaridids” by Jakob Vinther et al published in Science Advances.

The award-winning Everything Dinosaur website: Prehistoric Animal Models and Figures.

Researchers have named a new genus and species of rhabdodontomorph ornithopod from fossil material found in southeastern France. The dinosaur has been named Obelignathus septimanicus. The genus name was inspired by the French cartoon strip character Obélix from the Asterix the Gaul series, and the Latin word for jaw. Obélix is known for his exceptional strength and robust appearance. This is a reference to the unusually robustly built holotype dentary.

The researchers conclude that rhabdodontomorphs were more diverse in Europe than previously recognised.

A silhouette of the recently described 2025 ornithopod Obelignathus septimanicus from the Grès à Reptiles Formation in southern France. Scale bar equals 50 cm. Picture credit: Everything Dinosaur.

Picture credit: Everything Dinosaur

Obelignathus septimanicus

The origins and early diversification of the Ornithopoda is poorly understood.  For instance, the phylogeny of the Rhabdodontidae, a family of medium-sized ornithischian dinosaurs known solely from the Upper Cretaceous of Europe remains unresolved.  Current studies suggest eight or nine species, all of which are known from the upper Campanian to lower Maastrichtian of Europe.

Writing in the journal “Scientific Reports”, the researchers compiled a novel dataset made up from a morphological assessment and measurements of rhabdodontomorph dentaries.  The robust dentary (specimen number MDE D30) when first described in 1991 (Buffetaut and Le Loeuff), was thought to represent the genus Rhabdodon. However, it was sufficiently different to warrant its own species – Rhabdodon septimanicus. This latest analysis places R. septimanicus well outside the genus Rhabdodon and hence, a new genus has been erected.

Obelignathus is found to be a clear morphological outlier among European rhabdodontomorphs.  The research team members conclude that further large-scale studies are required to clarify the taxonomy of European rhabdodontomorphs.  These results indicate that this group of ornithopods exhibit greater diversity than currently recognised.  Several taxa appear to be coeval.

In common with most other rhabdodonts Obelignathus was relatively small. It is thought to have measured under three metres in length.

A New Herbivore from Late Cretaceous France

The find underscores southern France’s importance as a hotspot for Late Cretaceous dinosaur diversity. As fieldwork continues, Obelignathus offers a glimpse into the complex web of life that once thrived on the European archipelago.

Everything Dinosaur acknowledges the assistance of the open access scientific paper in the compilation of this article.

The scientific paper: “Exploring the diversity and disparity of rhabdodontomorph ornithopods from the Late Cretaceous European archipelago” by Łukasz Czepiński and Daniel Madzia published in Scientific Reports.

For models of ornithopods and other dinosaurs: Dinosaur Models.

A remarkable fossilised ichthyosaur flipper has provided new insights into the hunting behaviour of ancient marine reptiles.  The study, published in the journal “Nature” reveals that the giant ichthyosaur Temnodontosaurus trigonodon relied on stealth whilst hunting in the darkness. The metre-long front flipper was equipped with flow control structures that probably served to suppress self-generated noise as this megapredator hunted in dimly lit pelagic environments.

Life reconstruction of the giant Jurassic ichthyosaur Temnodontosaurus trigonodon hunting in the depths. The artwork highlights the winglike flipper, and the unusual structures observed in the fin. Picture credit: Joschua Knüppe.

Picture credit: Joschua Knüppe

Temnodontosaurus trigonodon Flipper Study

Temnodontosaurus was a large ichthyosaur. Size estimates vary, however, some individuals may have exceeded ten metres in length. Researchers have named several species within the genus. While fragmentary remains of unusual “fish lizards” had previously been excavated along the Dorset coast, it was the discovery of a metre-long skull by Joseph Anning at Lyme Regis in the autumn of 1811—followed by vertebrae and ribs found by 13-year-old Mary Anning in 1812 that prompted the first formal scientific study and description of an ichthyosaur. More than two centuries later, these enigmatic marine reptiles continue to yield unexpected insights and discoveries.

The fossilised flipper (specimen number SSN8DOR11) was collected from a temporary exposure of dark, laminated limestone (Lower Toarcian Posidonia Shale) of southwestern Germany. The partial front flipper preserves soft tissue structures and has been studied by an international team of researchers led by Dr Johan Lindgren from Lund University in Sweden.  The research was undertaken in collaboration with Dr Dean Lomax, an 1851 Research Fellow at the University of Bristol, who has been working on the fossil for about six years. Dr Lomax is one of the world’s leading ichthyosaur experts.

Dr Dean Lomax and Dr Johan Lindgren, together with fellow researcher Sven Sachs, examining one part of the Temnodontosaurus trigonodon flipper at Lund University, Sweden. Picture credit: Katrin Sachs.

Picture credit: Katrin Sachs

Dr Lindgren, who has pioneered research into marine reptile soft tissues commented:

“The wing-like shape of the flipper, together with the lack of bones in the distal end and distinctly serrated trailing edge collectively indicate that this massive animal had evolved means to minimise sound production during swimming. Accordingly, this ichthyosaur must have moved almost silently through the water, in a manner similar to how living owls—whose wing feathers also form a zigzag pattern—fly quietly when hunting at night. We have never seen such elaborate evolutionary adaptations in a marine animal before.”

The remarkable fossil flipper specimen (SSN8DOR11) shown left, under UV light (centre) and in a line drawing (right). Picture credit: Randolph G. De La Garza, Martin Jarenmark and Johan Lindgren.

Picture credit: Randolph G. De La Garza, Martin Jarenmark and Johan Lindgren

“Silent Swimming”

Although there are many exceptional ichthyosaur specimens with soft-tissue preservation, most soft tissues are associated with the fossilised remains of much smaller dolphin-sized species.  This is a remarkable discovery, it represents the first-ever soft tissues associated with a large-bodied ichthyosaur. In addition, the research team have identified unique structures never observed before in an aquatic animal.  The crenulated trailing edge of the wing-shaped flipper being reinforced by novel, mineralised, rod-like structures.  The researchers have named these structures “chondroderms”.

Dr Dean Lomax, who is also a palaeontologist at the University of Manchester, said:

“The first time I saw the specimen, I knew it was unique. Having examined thousands of ichthyosaurs, I had never seen anything quite like it. This discovery will revolutionise the way we look at and reconstruct ichthyosaurs (and possibly also other ancient marine reptiles) but specifically soft-tissue structures in prehistoric animals.”

Novel cartilaginous integumentary structures. To the left, light micrograph of the crenulated trailing edge in SSN8DOR11. Note that each serration is supported by a centrally located chondroderm. To the right, magnified image of a distal chondroderm. Picture credit: Randolph G. De La Garza, Martin Jarenmark and Johan Lindgren.

Picture credit: Randolph G. De La Garza, Martin Jarenmark and Johan Lindgren

The team postulate that this huge predator relied on underwater stealth, or “silent swimming” while hunting in the depths, in much the same way that owls as nocturnal predators have almost silent flight.

Eyes as Big as Footballs

Temnodontosaurus had the largest eyes of any vertebrate known.  The eye sockets of some specimens are more than twenty-five centimetres in diameter.  These huge eyes lend further support to the theory that Temnodontosaurus hunted under low-light conditions, either at night or in deep waters.

Spectacular, 183-million-year-old soft-tissue fossil (SSN8DOR11; Paläontologisches Museum Nierstein, Nierstein, Germany): an isolated wing-like front flipper of the giant predatory ichthyosaur Temnodontosaurus (T. trigonodon). Photograph (left) and shown under UV light (centre). Line drawing (right) providing a representation of the metre-long flipper. Picture credit: Randolph G. De La Garza, Martin Jarenmark and Johan Lindgren.

Picture credit: Randolph G. De La Garza, Martin Jarenmark and Johan Lindgren

The remarkable fossilised flipper was discovered by fossil collector Georg Göltz, a co-author on the new study. Amazingly, Georg made the find entirely by chance whilst looking for fossils at a temporary exposure at a road cutting in the municipality of Dotternhausen, Germany. The fossil consists of both the part and counterpart (opposing sides) of almost an entire front flipper.  Georg continued to look for more remains, but no other fossil material was found.

As the proximal part of the fin is absent, it has been speculated that the flipper could have been ripped off by an even larger ichthyosaur.  The specimen was shown to palaeontologist and study co-author Sven Sachs (Natural History Museum, Bielefeld).  Dr Sachs immediately recognised the rarity of the find.

A multidisciplinary research team employed a variety of sensitive imaging, elemental and molecular analyses to examine the unique preserved structures. This involved high-end techniques such as synchrotron radiation-based X-ray microtomography at the Swiss Light Source SLS at PSI and Diamond Light Source, time-of-flight secondary ion mass spectrometry and infrared microspectroscopy, along with the reconstruction of a virtual model using computational fluid dynamics.

To recreate the stealth hunting behaviours of a marine reptile that lived more than 180 million years ago is remarkable. Furthermore, by studying the fin morphology, scientists could find ways of reducing our impact on modern marine soundscapes.

Going Back to Mary Anning

For Dr Dean Lomax, this astonishing study harks all the way to back to the days of pioneering palaeontologist Mary Anning and her older brother Joseph.

He stated:

“In a weird way, I feel that there is a wonderful full-circle moment that goes back to Mary Anning showcasing that even after two hundred years, we are still uncovering exciting and surprising finds that link back to her initial discoveries.”

Dr Lomax added:

“The fossil provides new information on the flipper soft tissues of this enormous leviathan. It has structures never seen in any animal, and reveals a unique hunting strategy thus providing evidence of its behaviour, all combined with the fact that its noise-reducing features may even help us to reduce human-made noise pollution. Although I might be a little bias, in my opinion, this represents one of the greatest fossil discoveries ever made.”

Everything Dinosaur acknowledges the assistance of a media release from the University of Bristol in the compilation of this article.

The scientific paper: “Adaptations for stealth in the wing-like flippers of a large ichthyosaur” by Johan Lindgren, Dean R. Lomax, Robert-Zoltán Szász, Miguel Marx, Johan Revstedt, Georg Göltz, Sven Sachs, Randolph G. De La Garza, Miriam Heingård, Martin Jarenmark, Kristina Ydström, Peter Sjövall, Frank Osbæck, Stephen A. Hall, Michiel Op de Beeck, Mats E. Eriksson, Carl Alwmark, Federica Marone, Alexander Liptak, Robert Atwood, Genoveva Burca, Per Uvdal, Per Persson and Dan-Eric Nilsson published in the journal Nature.

Visit the website of the award-winning palaeontologist and author Dr Dean Lomax: Palaeontologist Dr Dean Lomax.

Researchers have described a new species of Late Triassic pterosaur.  It has been named Eotephradactylus mcintireae and it represents North America’s oldest known flying reptile. A field team led by the Smithsonian National Museum of Natural History and Columbia College Chicago discovered the fossil remains. They were exploring outcrops of the Owl Rock Member of the Chinle Formation in north-eastern Arizona. They come from an unusual bonebed that preserves the remains of a variety of Late Triassic vertebrates. The assemblage provides evidence of animals that persisted into the Jurassic co-existing with archaic lineages such as phytosaurs, armoured aetosauriforms and ancient temnospondyls.

The bonebed provides a snapshot of an ecosystem prior to the End Triassic Mass Extinction event.

North America’s oldest pterosaur described to date Eotephradactylus mcintireae disturbs some frogs whilst catching a ray-finned fish. An early tortoise is seen in the background and the bones of an armoured crocodilian are visible. Picture credit: Brian Engh.

Picture credit: Brian Engh

Eotephradactylus mcintireae – North America’s Oldest Pterosaur

The study has been published in the “Proceedings of the National Academy of Sciences.”  The pterosaur fossils identified from the bonebed include a partial left mandible (lower jaw), isolated teeth and a phalanx (digit bone).  The fossils are approximately 209 million years old (Norian faunal stage of the Late Triassic).

Corresponding author Ben Kligman (Peter Buck Postdoctoral Fellow at the Smithsonian’s National Museum of Natural History), stated:

“The site captures the transition to more modern terrestrial vertebrate communities where we start seeing groups that thrive later in the Mesozoic living alongside these older animals that don’t make it past the Triassic. Fossil beds like these enable us to establish that all of these animals actually lived together.”

Kay Behrensmeyer (left), the curator of vertebrate palaeontology at the Smithsonian’s National Museum of Natural History. Robin Whatley (right), professor and associate dean at Columbia College Chicago in the Petrified Forest National Park digging for fossils in a quarry in 2023. Picture credit: Ben Kligman, (Smithsonian).

Picture credit: Ben Kligman (Smithsonian)

“Ash-winged Dawn Goddess”

The genus name translates as “ash-winged dawn goddess”, it references the location’s volcanic ash layers and the pterosaur’s position as a basal member of the Pterosauria family tree. The species name honours preparator Suzanne McIntire, who discovered the fossil material in 2013 when preparing a block from the site.

The pterosaur fossil was unearthed by preparator Suzanne McIntire, a volunteer at the museum’s FossiLab for eighteen years. Picture credit: Bill King.

Picture credit: Bill King

The Owl Rock Member

The Owl Rock Member strata are some of the least explored parts of the Petrified Forest National Park.  The extensive volcanic ash layers in the quarry, permitted the scientists to calculate the age of the bonebed.  It represents the remains of an ecosystem that existed around 209 million years ago (Norian faunal stage of the Late Triassic).  These are some of the geologically youngest sediments in the Petrified Forest National Park.

In the Late Triassic, Arizona was positioned in the central part of the supercontinent Pangaea. The area was just north of the equator. It was a low-lying, semi-arid environment criss-crossed with small river channels and prone to seasonal floods. The bonebed likely preserves the remains of a community of animals that were caught up in a flash flood.

The bonebed is rich vertebrate fossils. So many fossils were found that excavating them in the field was impossible. Field team members took large chunks of rock, entombed in plaster jackets back to the preparation laboratory at the Smithsonian National Museum of Natural History. Volunteers spent thousands of hours carefully excavating the fossil material.

Ben Kligman (right), a Peter Buck Postdoctoral Fellow at the Smithsonian’s National Museum of Natural History with several of the museum’s FossiLab volunteers who contributed to the study of the fossils from the bonebed. Richard Cline (far left), Hillary Cochard, James Morrison and far right Lynn Sharp. Picture credit: Abby Telfer, (Smithsonian).

Picture credit: Abby Telfer (Smithsonian)

A Diverse Community of the Familiar and Not So Familiar

The gull-sized pterosaur Eotephradactylus mcintireae shared its home with a huge variety of other vertebrates. Some of these animals would be familiar to us, others represent taxa that did not survive into the Jurassic. For example, coelacanths swam in the rivers along with freshwater sharks.  In addition, frogs were present and an ancient tortoise.

The tortoise fossils are of particular interest. They are some of the world’s oldest tortoise remains. The tortoise had a robust shell, with protruding spikes for additional protection. Stem members of the Testudinata clade (reptiles with a true carapace and plastron shell) are known from the Late Triassic. For instance, the genus Proterochersis was present in Germany and Poland and is roughly contemporaneous with the Owl Rock Member biota.

Ben Kligman commented:

“This suggests that turtles rapidly dispersed across Pangaea, which is surprising for an animal that is not very large and is likely walking at a slow pace.”

Animals that are familiar to us today coexisted with animals that were very different to modern faunas.  For example, whilst relatives of New Zealand’s tuatara were present (rhynchocephalians), there were also giant, ancient amphibians, aetosaurs and phytosaurs.

Over 1,200 Fossils Including the Pterosaur Eotephradactylus mcintireae

The team has uncovered more than 1,200 individual fossils. They include bones, teeth, fish scales and coprolites. Sixteen different types of vertebrate were identified from this single bonebed indicating a diverse ecosystem.  Amongst these fossils were the remains of North America’s oldest known pterosaur Eotephradactylus mcintireae.  Its discovery motivates research teams to continue to explore the remote and difficult to access Owl Rock Member exposures.

Suzanne McIntire reflecting on the significance of her discovery said:

“What was exciting about uncovering this specimen was that the teeth were still in the bone, so I knew the animal would be much easier to identify.”

The teeth provide important clues to the pterosaur’s diet; the worn-down crowns suggest it was durophagous, adapted for consuming hard-shelled prey.  The researchers conclude that this flying reptile fed on the site’s fish. Ray-finned fish are known from this locality (actinopterygians).  Many had armour-like scales, and a diet of these types of fish would have resulted in extensive tooth wear.

Everything Dinosaur acknowledges the assistance of a media release by the Smithsonian National Museum of Natural History in the compilation of this article.

The scientific paper: “Unusual bone bed reveals a vertebrate community with pterosaurs and turtles in equatorial Pangaea before the end-Triassic extinction” by Ben T. Kligman, Robin L. Whatley, Jahandar Ramezani, Adam D. Marsh, Tyler R. Lyson, Adam J. Fitch, William G. Parker and Anna K. Behrensmeyer published in the Proceedings of the National Academy of Sciences.

The award-winning Everything Dinosaur website: Pterosaur and Prehistoric Animal Models.