In 1984, a field team led by the renowned Argentinian palaeontologist José Bonaparte uncovered the fossilised remains of a theropod dinosaur in Chubut Province (Patagonia). The articulated fossil remains included most of the front-portion of the skeleton and although some of the bones had been deformed and distorted due to taphonomic processes, skin impressions of parts of the right side of the animal had been preserved. Skin impressions associated with the head of this dinosaur were also present, but these were not recognised during laboratory preparation and sadly they were lost as the skull fossils were cleaned and prepared.

A year later, Carnotaurus (C. sastrei) was formally named and described. Remarkably, despite the Carnotaurus skin impressions being the most completely preserved of any theropod, no detailed study of the skin had been undertaken.

All that changed this week with the publication of a scientific paper in the journal “Cretaceous Research”.

Not a Feather to be Found

Palaeontologist Dr Christophe Hendrickx from the Unidad Ejecutora Lillo in San Miguel de Tucumán (Argentina), worked with Dr Phil Bell from the University of New England (New South Wales, Australia), an expert in dinosaur integumentary coverings. Whilst the skin impressions only cover part of the body, (the largest skin impression is associated from the base of the tail), the scientists were able to determine that the skin covering consisted of a diverse range of scales and bumps of different shapes and sizes.

No evidence for any bristle-like structures or feathers could be found.

Dr Hendrickx remarked:

“By looking at the skin from the shoulders, belly and tail regions, we discovered that the skin of this dinosaur was more diverse than previously thought, consisting of large and randomly distributed conical studs surrounded by a network of small elongated, diamond-shaped or sub-circular scales.”

Have Carnotaurus Model Makers Got it All Wrong?

Contrary to previous interpretations and the attempts of model makers to depict Carnotaurus, the feature scales are randomly distributed and neither form discrete rows nor show progressive variations in their size along parts of the body. All those illustrations and replicas of Carnotaurus with a neat row of spines running down its back are not accurate according to the conclusions drawn from this research.

Picture credit: Everything Dinosaur

The picture above shows a Carnotaurus replica in the Nanmu Studio Jurassic Series range.

To view this range: Nanmu Studio Dinosaur Models.

Carnotaurus Skin – Reminiscent of a Thorny Devil

The composition of the skin and the morphology of the scales reminded the researchers of the integument of the Australian lizard the Thorny Devil (Moloch horridus). This small, spiny lizard which is relatively common in the deserts of western and central Australia, uses its spines primarily for defence. It would be difficult for any would-be predator to swallow it. Grooves between the spines allow the lizard to channel water to its mouth, a useful adaptation when living in an environment with infrequent rain.

At around 8 metres in length and since Carnotaurus is regarded as the apex predator in its environment, it is unlikely that the lumps and bumps on the skin were primarily for self-defence, but protection from intraspecific combat cannot be ruled out. However, recent studies have shown that Carnotaurus was a strong runner. If this large dinosaur had a very active lifestyle, then helping to regulate body temperature and permit heat-loss would have been very important.

Playing a Role in Thermoregulation

The researchers speculate that the skin may have played a vital role in thermoregulation, a role consistent with integument function in living mammals and reptiles.

The scientific paper: “The scaly skin of the abelisaurid Carnotaurus sastrei (Theropoda: Ceratosauria) from the Upper Cretaceous of Patagonia” by Christophe Hendrickx and Phil R. Bell published in Cretaceous Research.

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A new species of carcharodontosaurid has been named from a single fragment of upper jawbone found in Uzbekistan. The dinosaur has been named Ulughbegsaurus uzbekistanensis and it was probably the apex predator in the ecosystem suggesting that carcharodontosaurids remained the dominant predators relative to tyrannosauroids, at least in Asia until around 90 million years ago.

The First Late Cretaceous Carcharodontosaurian from Central Asia

Ulughbegsaurus has been named based on partial maxilla found in strata associated with the Bissekty Formation of the Kyzylkum Desert (Uzbekistan). Several other predatory theropods have been described from fossils found in this formation, but all of them are considerably smaller. The tyrannosauroid Timurlengia euotica was coeval, but much smaller than Ulughbegsaurus providing further support for the idea that carcharodontosaurians were the dominant, apex predators in Laurasia until their extinction some 20 million years prior to the end of the Cretaceous, from which point onwards it was the tyrannosauroids that took over this niche in most Laurasian ecosystems.

Estimating Size from a Single Fragment of Bone

Palaeontologists can use the size of the tooth row in the maxilla to help them estimate the body size of theropod dinosaurs. Studies of carcharodontosaurids and tyrannosaurids have demonstrated that the length of the tooth row in the maxilla is isometrically correlated with femur length, which is very helpful, as the length of the thigh bone is widely used to help calculate body mass. Based on this data, the authors of this paper, calculate that the Ulughbegsaurus specimen was at least 7 metres long and over a tonne in weight.

The researchers, which included corresponding author Kohei Tanaka (University of Tsukuba, Japan) and Darla Zelenitsky (University of Calgary, Canada), conclude that the individual represented by the single bone was probably 7.5 to 8 metres in length.

Ulughbegsaurus uzbekistanensis was much bigger than any other theropod known from this region. The tyrannosauroid Timurlengia was approximately 3-4 metres long and around 8 times lighter. This suggests that Timurlengia was a secondary predator along with an as yet, unnamed large dromaeosaurid, whilst Ulughbegsaurus occupied the niche of apex predator. The discovery of Ulughbegsaurus records the geologically latest stratigraphic co-occurrence of carcharodontosaurid and tyrannosauroid dinosaurs from Laurasia and evidence indicates carcharodontosaurians remained the dominant predators relative to tyrannosauroids, at least in Asia, as late as the Turonian faunal stage of the Cretaceous.

Tyrannosauroids Kept in Check by Carcharodontosaurians

For much of the Cretaceous allosauroids (part of the Carnosauria clade), including carcharodontosaurians were the largest terrestrial predators on Earth. It was only after their extinction that tyrannosauroids (members of another theropod clade, the Coelurosauria), became much larger and occupied the role of apex predators in most ecosystems across Laurasia.

Evidence of larger tyrannosauroids is not known until the Campanian of North America, some 7 million years after Ulughbegsaurus and Timurlengia lived. Palaeontologists remain uncertain as to the dynamics of apex predator evolution amongst the Theropoda as the fossil record from 90 to 83 million years ago (Coniacian-Santonian) is extremely poor.

Honouring a Sultan of the Timurid Empire

Ulughbegsaurus uzbekistanensis (pronounced Ul-lug-bey-sore-us uz-bek-ee-stan-en-sis), was named in honour of Ulugh Beg, a sultan and polymath of the Timurid Empire in the fifteenth-century. The species or trivial name honours the country of Uzbekistan.

A Significant Fossil Discovery

Although Ulughbegsaurus has been described from a single bone, its discovery is very significant. U. uzbekistanensis represents the first definitive fossil evidence of carcharodontosaurians from Central Asia. It fills a geographic gap in the clade between Europe and East Asia and shows that carcharodontosaurians were widespread across Asia.

To read Everything Dinosaur’s 2016 article about the discovery of Timulengia euotica: Fossil Study Shows How Tyrannosaurs Got Big.

To read about the diminutive tyrannosauroid Moros intrepidus that co-existed with the much larger allosauroid Siats meekerorum: Fleet-footed Tyrannosaur Leaps 70 million-year Gap.

The scientific paper: “A new carcharodontosaurian theropod dinosaur occupies apex predator niche in the early Late Cretaceous of Uzbekistan” by Kohei Tanaka, Otabek Ulugbek Ogli Anvarov, Darla K. Zelenitsky, Akhmadjon Shayakubovich Ahmedshaev and Yoshitsugu Kobayashi published in Royal Society Open Science.

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

Researchers from the Royal Ontario Museum (Canada), have announced the discovery of a new species of armoured arthropod from the Burgess Shale of British Columbia. A study looking at 12 fossil specimens collected from Marble Canyon and Tokumm Creek in the Kootenay National Park (British Columbia), has been published this week by Royal Society Open Science. The arthropod has been named Titanokorys gainesi and at around 50 cm in length, it is a giant by Cambrian biota standards.

The authors of the scientific paper, Dr Jean-Barnard Caron of the Royal Ontario Museum, an expert on Burgess Shale fauna and PhD student Joe Moysiuk, classify Titanokorys as a member of the Radiodonta, a stem group of the Arthropoda. Radiodonts were extremely diverse and geographically widespread during the Late Cambrian and many of them were giants when compared to other animals alive during this time in Earth’s history. Perhaps the most famous radiodont is the taxon Anomalocaris, regarded by many palaeontologists as the world’s first super-predator.

The CollectA Anomalocaris model. A fantastic replica of an early apex predator. The CollectA Anomalocaris (Other Prehistoric Animal Models).

The picture (above) shows a model of an Anomalocaris.  This figure is part of the CollectA Prehistoric Life model range.

To view this range: CollectA Prehistoric Life.

Living on the Seabed – A Benthic Existence

Radiodonts are characterised by their compound eyes, disc-shaped mouthparts and paired frontal appendages, which in the case of Titanokorys consist of comb-like structures which may have been used to sift through mud in search of prey. The broad, flattened carapace of Titanokorys supports the idea that it was benthic – living on the seabed.

Dr Caron stated:

“The sheer size of this animal is absolutely mind-boggling, this is one of the biggest animals from the Cambrian period ever found.”

Coeval with Cambroraster falcatus

The bedding planes that provided the Titanokorys fossil material have also revealed an abundance of the smaller, but closely related Cambroraster falcatus, which was named and described by Caron and Moysiuk in 2019. Cambroraster was named after the Millennium Falcon from Star Wars franchise, as its carapace resembled the shape of this iconic spaceship. The co-occurrence of these two species on the same bedding planes highlights potential competition for benthic resources and the high diversity of large predators sustained by Cambrian communities.

To read about the discovery of Cambroraster falcatus: Prehistoric Predator with a Mouth Shaped Like a Pineapple Ring.

Why some radiodonts evolved such a bewildering array of head carapace shapes and sizes is still poorly understood and was likely driven by a variety of factors.

Dr Caron added:

“These enigmatic animals certainly had a big impact on Cambrian seafloor ecosystems. Their limbs at the front looked like multiple stacked rakes and would have been very efficient at bringing anything they captured in their tiny spines towards the mouth. The huge dorsal carapace might have functioned like a plough.”

Honouring Professor Robert Gaines

The genus name is derived from the Greek Titans, powerful gods of huge size and from the Greek “Korys” for helmet. The species or trivial name honours Professor Robert Gaines who was instrumental in the co-discovery of the Marble Canyon fossil deposit, where some of the Titanokorys specimens were found.

The scientific paper: “A giant nektobenthic radiodont from the Burgess Shale and the significance of hurdiid carapace diversity” by J.B. Caron and J. Moysiuk published by Royal Society Open Science.

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A superbly preserved and almost complete specimen of the pterosaur Tupandactylus navigans has enabled researchers to get a better understanding of the body of this Brazilian flying reptile. The skull too, with its exquisite preservation has provided new data on the amazing super-sized sagittal crest associated with this genus.

Confiscated in a Police Raid

The fossil specimen (number GP/2E 9266), was confiscated in a police raid at Santos Harbour, São Paulo State (Brazil), along with several other beautifully well-preserved fossils. Unfortunately, illegal fossil collection and sale of specimens on the black market is an increasing problem in Brazil. However, the successful raid prevented this hugely significant pterosaur fossil from ending up in the hands of a private collector.

The specimen is now housed at the Universidade de São Paulo (Brazil) and most of the researchers involved in the scientific paper are also based in Brazil, except for Octávio Mateus (Museu da Lourinhã, Portugal). Writing in the on-line, open access journal PLOS One, the scientists conclude that this specimen is the best-preserved tapejarid skeleton discovered to date. Their analysis has shed new light on the anatomy of this Tapejaridae family.

CT Scans

The pterosaur fossil is comprised of six limestone slabs. As this fossil was collected illegally, its provenance is unknown. However, by studying the yellow-stained, laminated limestone matrix, the research team were able to confidently assign the fossil to the Aptian-aged Crato Formation. The fossil took a trip to the local hospital in São Paulo, to enable CT scans to be undertaken. The subsequent three-dimensional models generated enabled the scientists to reconstruct the body of Tupandactylus for the first time. Prior to the discovery of this fossil, most Brazilian tapejarids had been described based on isolated skull bones.

Tupandactylus and a Remarkable Head Crest

Soft tissue from the huge crest on the top of the head, extends to more than five times the actual height of the skull. Analysis of the crest enabled the research team to confirm differences between Tupandactylus navigans and the closely related T. imperator. Specimens of T. navigans tend to be smaller than T. imperator and it had been speculated that just one species was represented with the differences between T. imperator and T. navigans being explained by sexual dimorphism. Thanks to the exquisite preservation of this fossil specimen the research team were able to identify several anatomical traits that support the idea that Tupandactylus navigans and Tupandactylus imperator are indeed separate species.

Different colour patterns seen in the sagittal crest do not represent fossilisation of any colour patterning but may have occurred due to oxidation of the material. However, a more detailed analysis of the soft tissues associated with GP/2E 9266 is currently being undertaken.

Tupandactylus navigans

Tupandactylus navigans was named and scientifically described in 2003 based on a study of two fossil skulls (Frey, Martill and Buchy). At the time, it was postulated that the huge head crest could have been used as a sail to help with flight stability and assist with aerial propulsion. For this structure to work in this way, the neckbones would have had to be robust, relatively short and supported by powerful tendons to combat stresses imposed on the neck. With an almost complete, articulated skeleton to study, the scientists led by Victor Beccari (Universidade de São Paulo), discovered that this pterosaur had a long neck, long limbs and relatively short wings (estimated wingspan 2.7 metres).

The enormous crest probably did not play a role in aiding powered flight. Such a huge structure may well have hindered this pterosaur’s aerial abilities.

A spokesperson from Everything Dinosaur commented:

“The sagittal crest of Tupandactylus may have evolved due to sexual selection pressure, females showing a bias towards males with larger crests. The crest may have been used for display or to denote maturity, status or fitness for breeding. Many living birds sport crests, wattles and other structures, perhaps adult T. navigans sacrificed some aerial ability by growing enormous crests in a bid to attract mates.”

For models and replicas of pterosaurs and other prehistoric creatures: Prehistoric Animal Models.

Spotting a Notarium

The front five dorsal vertebrae form a notarium which helps brace the chest and counter the stresses on the torso created by the flapping of the wings. This structure is found in living birds and some types of pterosaur but this is the first time, as far as Everything Dinosaur team members are aware, that a notarium has been identified in a tapejarid.

he scientific paper: “Osteology of an exceptionally well-preserved tapejarid skeleton from Brazil: Revealing the anatomy of a curious pterodactyloid clade” by Victor Beccari, Felipe Lima Pinheiro, Ivan Nunes, Luiz Eduardo Anelli, Octávio Mateus and Fabiana Rodrigues Costa published in PLOS One.

Researchers writing in the on-line, open-access journal PLOS One have described the most complete plesiosaur skull found to date from sub-Saharan Africa. The skull and post cranial fossil material reported upon comes from the Upper Cretaceous Mocuio Formation exposures located at Bentiaba in the Namibe Province of Angola.

The fossils represent another specimen of the elasmosaurid Cardiocorax (C. mukulu) one of just six valid plesiosaurian taxa known from the whole of Africa. This plesiosaur swam in a shallow, tropical sea and the fossils are around 71.5 million years old.

Cardiocorax mukulu

The specimen was found in sandstone deposits, stratigraphically just 3 metres above where the holotype material for this taxon was found. The researchers who studied the fossils, which included Octávio Mateus (Museu da Lourinhã, Portugal) and lead author Miguel Marx from the Southern Methodist University (Dallas, Texas), concluded that the fossils represented C. mukulu as the overlapping skeletal material of this specimen – MGUAN PA278 with that of the holotype were virtually identical.

Picture credit: Everything Dinosaur

This elasmosaurid had been originally named and described back in 2015. The genus name comes from the Greek “kardia” which means heart and coracoid, a paired bone that forms part of the shoulder in most vertebrates. This is a reference to the unique heart-shaped fenestra (hole) which occurs between the coracoid bones, a characteristic that is unique to this genus. The specific or trivial name comes from the local Angolan Bantu dialect and means “ancestor).

High Resolution CT Scans Reveal Details of Plesiosaur Skull

The exquisite, three-dimensional preservation of the skull material provided the scientists with an opportunity to examine in close detail the morphology of the skull. The delicate fossil could have been damaged during further preparation, so the specimen was subjected to high resolution CT scans at the University of Texas High-Resolution X-ray CT Facility this permitted the anatomy of the skull to be revealed.

The holotype fossils, which were used to name and describe this elasmosaurid back in 2015 lacked skull bones. The discovery of an almost complete, three-dimensional skull of Cardiocorax enabled the research team to conduct an elaborate series of phylogenetic assessments to assess where within the Plesiosauria Cardiocorax should be placed.

Most of these analyses suggest an early-branching or intermediate position for Cardiocorax mukulu within the Elasmosauridae family. Elasmosaurines have elongated neck bones (cervical vertebrae), this anatomical characteristic is absent in Cardiocorax mukulu which suggests that it was a relic of an older, less derived lineage of elasmosaurids. These results indicate that several different types of elasmosaurid persisted into the Maastrichtian faunal stage of the Cretaceous.

The scientific paper: “The cranial anatomy and relationships of Cardiocorax mukulu (Plesiosauria: Elasmosauridae) from Bentiaba, Angola” by Miguel P. Marx, Octávio Mateus, Michael J. Polcyn, Anne S. Schulp, A. Olímpio Gonçalves and Louis L. Jacobs published in PLOS One.

The Everything Dinosaur website: Prehistoric Animal Figures and Toys.

In 2017, Everything Dinosaur reported on the discovery of an extensive pterosaur nesting site that had been found south of the Tian Shan Mountains in Xinjiang (Xinjiang Uygur Autonomous Region), in north-western China. The bonebeds represented colonial nesting sites of Hamipterus tianshanensis. It was a flying reptile that had been formally named and described back in 2014.

To read Everything Dinosaur’s 2017 article about the Hamipterus nesting sites: Hamipterus Nesting Ground Discovery.

New Sauropods Described

Fieldwork in this area has revealed the presence of dinosaurs and a paper has been published this week describing sauropod fossils. These are the first dinosaurs to be identified from this part of China. Two of the fossil specimens have led to the erection of new sauropod species Silutitan sinensis and Hamititan xinjiangensis. Four incomplete bones from the sacrum along with associated sacral ribs representing another sauropod specimen have also been found, but as yet, no genus for these fossils has been assigned.

Silutitan sinensis

Described from a series of six articulated cervical vertebrae (neck bones), with associated cervical ribs, Silutitan sinensis (which translates as Chinese Silk Road Titan), has been assigned to the Euhelopodidae family and is thought to have been closely related to Euhelopus (E.zdanskyi). At an estimated twenty metres plus, it is the larger of the two sauropods to be described in the scientific paper.

Hamititan xinjiangensis

Although contemporaneous with Silutitan (Lower Cretaceous, Shengjinkou Formation of the Tugulu Group). Hamititan was not closely related. Cladistic analysis based on the tail bones (caudal vertebrae) from which this dinosaur was described, suggests that Hamititan is a titanosaur (a lithostrotian titanosaur), as such, it seems more closely related to South American titanosaurs such as Epachthosaurus and Notocolossus.

The genus name translates as Hami City Titan, whilst the species name honours the Xinjiang Uygur Autonomous Region of China.

A single theropod tooth (below, picture F), indicates the presence of theropod dinosaurs.

No Name for the Third Specimen

A third specimen consisting of four sacrum bones and associated sacral ribs has not been assigned to a species. However, the researchers, writing in the academic journal Scientific Reports conclude that not only do these fossils represent the first dinosaurs to be described from the Shengjinkou Formation but they also help to support the hypothesis that Asia was home to a great variety of different types of sauropod during the Early Cretaceous.

The scientific paper: “The first dinosaurs from the Early Cretaceous Hami Pterosaur Fauna, China” by Xiaolin Wang, Kamila L. N. Bandeira, Rui Qiu, Shunxing Jiang, Xin Cheng, Yingxia Ma and Alexander W. A. Kellner published in Scientific Reports.

The Everything Dinosaur website: Dinosaur Models and Toys.