feather all the dinosaurs – Nix Illustration

It Came From The Wastebasket #10: Struggling With Stegoceras

First described and named in the early 1900s, Stegoceras validum was a dog-sized small pachycephalosaur that lived in Alberta, Canada, during the Late Cretaceous (~77-74 million years ago).

Initially just known from its skull domes, it was one of the first pachycephalosaurs to be discovered and was very poorly understood until more complete remains were found in the 1920s. Then it spent a couple of decades being mixed up with Troodon due to similarities in tooth shape, until the discovery of Pachycephalosaurus led to pachycephalosaurs finally being recognized as a distinct group of ornithischian dinosaurs in the 1940s.

An illustration of Stegoceras, an extinct pachycephalosaur. It's a small bipedal dinosaur with tiny arms, bird-like legs, a speculative coat of fluffy protofeathers over most of its body, and a long tapering tail with speculative bristly quills. It has a large bony dome on its forehead, rimmed with short spikes, and a short snout with a stubby beak. It's mainly colored white and grey, with brighter red and yellow markings on its face and red towards the tip of its tail. — *Stegoceras validum*

For much of the 20^th century Stegoceras was treated as a wastebasket taxon for any small-to-mid-sized North American (and one Asian) pachycephalosaur, and multiple different species were named based on what were often rather dubious fragmentary fossils. But towards the start of the 21^st century this mess did start getting cleaned up, merging some dubious species into the original Stegoceras validum, and moving others to separate genera like Sphaerotholus, Colepiocephale, Hanssuesia, and Sinocephale.

By the early 2000s just the Canadian Stegoceras validum remained – but then in 2011 the new species Stegoceras novomexicanum was named based on specimens from New Mexico, USA. The validity of this second species has been debated, since the fossils are juveniles and might instead belong to Stegoceras validum or another genus like Sphaerotholus, but if it is some sort of Stegoceras then it significantly re-extends the known geographic range of this little pachycephalosaur.

Retro vs Modern #21: Deinonychus antirrhopus

Deinonychus antirrhopus was one of the most significant dinosaur discoveries of the 20^th century, kicking off the Dinosaur Renaissance and the recognition of the evolutionary link between maniraptoran theropods and modern birds.

1960s

The first remains of this species were discovered in North America in the 1930s, but at the time the fossils weren’t officially described or named. It wasn’t until the 1960s that more specimens were found in Montana, representing at least three preserved individuals, and paleontologist John Ostrom recognized that there was something very special about this dinosaur.

In contrast to the prevailing view at the time that theropods were all upright tail-dragging “sluggish lizards” this was clearly a highly specialized and active predator, with a huge sickle-shaped claw on each foot and a long stiff tail for balance – inspiring its scientific name’s meaning of “counterbalancing terrible claw”.

And while the very first reconstruction of Deinonychus might seem retro to modern eyes, at the time it was revolutionary and it went on to become an iconic representation of the species for the next couple of decades. Drawn by Robert Bakker, who was Ostrom’s student at the time, it depicted a lizard-like creature with its body held in a horizontal pose and its tail held out straight behind it. Its head was portrayed as more domed than we now know Deinonychus’ skull to have been, and its neck was up in an alert posture while the animal ran at full sprint, with its sickle-claws held up away from the ground to keep them sharp.

A few years later further discoveries showed a highly bird-like pelvis and hands very similar to those of Archaeopteryx, triggering the Dinosaur Renaissance reinterpretation of dinosaurs as active warm-blooded animals, and the revival of the 19^th century idea that they were the ancestors of birds.

1980s-1990s

As the “birds are dinosaurs” idea began to gain acceptance with increasing amounts of anatomical evidence, some paleontologists in the 1980s began to also suspect that highly bird-like dromaeosaurids like Deinonychus might have also been feathered. Some reconstructions during this time showed this to varying degrees, particularly those drawn by Bakker and by Gregory Paul – but it didn’t really catch on more widely at first, for one very big reason:

Jurassic Park happened.

Dromaeosaurs hadn’t been well-known dinosaurs to the general public before that point, but the 1993 JP “raptors” were an instant hit in pop culture. Physically based much more on Deinonychus than on Velociraptor, and exemplifying the renaissance view of dinosaurs in major media for the first time, the movie’s fully scaled and oversized version of these animals dominated popular depictions for years afterwards. Even the most rigorous and anatomically accurate artwork showcasing their bird-like features still usually kept them completely naked to retain that familiar reptilian appearance.

Most 1990s attempts at any feathering tended towards being as sparse as possible – often along with the shrinkwrapping typical for the era – at best being decent for the time but what we’d now deride as “half-assed”, and at worst being “a few token quills on the back of the head”.

Deinonychus fossils found in association with Tenontosaurus were also interpreted as being evidence of cooperative pack hunting behavior during this time, and it became a common paleoart meme to depict the large herbivore being constantly swarmed by ravenous raptors.

2020s

The mid-1990s discovery of fully-feathered dinosaurs like Sinosauropteryx in China, followed a few years later by raptors with wing-feathered arms like Sinornithosaurus, gradually began to put the fluff back onto dinosaurs like Deinonychus.

(…At least in reasonably scientific paleoart. The much much stronger and ongoing resistance from popular culture is far too big a subject to get into here. But maybe, just maybe, we’re finally hitting a turning point there?)

Early attempts at properly feathering dinosaurs were a bit awkward, usually looking rather like a bunch of scruffy greasy hair glued onto a scaly raptor, a dinosaur wearing fuzzy pajamas, or like the old “bird-lizard” depictions of Archaeopteryx. Even into the early 2010s some paleoart memes were still common in depictions of dromaeosaurs, but increasingly better understanding of their anatomy and plumage arrangements over the last decade or so has brought us to a much more birdlike interpretation of these animals – with paleoartists like Emily Willoughby being especially influential in popularizing the modern view of dinosaurs like Deinonychus.

We now know Deinonychus lived during the Early Cretaceous, about 115-108 million years ago, in what is now the Mountain West and South Central United States. Up to around 3.4m long (11′), it stood about 1m tall (3’3″), similar in size to a large dog.

It had blade-like teeth in its jaws, and forward-facing eyes with stereoscopic vision. Its three-fingered arms would have been covered by wing-like feathers, and its tail probably had feathers all the way along its length and was stiffened but not totally inflexible.

It may have used the sickle-claws on its feet to pin down struggling prey, eating it alive while flapping its wings and waving its tail for balance. And while often depicted as an extremely fast-runner, its leg proportions and foot anatomy suggest it was actually built more for walking and had an especially strong grip strength in its feet, trading speed for power and probably being more of an ambush predator – often being compared to a “giant ground-hawk”.

Pack hunting has been called into question recently, too, arguing that the Tenontosaurus sites may actually represent crocodile-like or Komodo dragon-like behavior with mobs of scavenging individuals congregating at a carcass. But other evidence from trackways and Utahraptor does offer potential support for pack behavior in raptors, so it’s still open to interpretation.

Retro vs Modern #10: Plateosaurus trossingensis

First discovered in southeast Germany in the 1930s, Plateosaurus was only the fifth non-avian dinosaur known to science – but its fossils were fragmentary and poorly understood until the early 20^th century, when large bonebeds full of much better specimens began to be excavated.

1910s-1970s

Between the 1910s and 1930s around 80 near-complete skeletons of Plateosaurus were found in two German quarries, quickly making it one of the most abundant and best known dinosaur species of the time. Although it had previously been classified as a theropod dinosaur, in the 1920s the more complete material allowed it to be properly identified as a “prosauropod“, an early herbivorous relative of the giant sauropods.

Like many bipedal dinosaurs during this period Plateosaurus was generally interpreted as having an upright kangaroo-like posture with a dragging tail – although some paleontologists were arguing for it having a sprawling quadrupedal lizard-like stance as late as the mid-1930s.

1980s

Unfortunately much of the German fossil material was destroyed during World War II bombing raids, and interest in Plateosaurus didn’t pick up again until the time of the Dinosaur Renaissance when a third major fossil site was discovered in Switzerland during the 1970s.

Plateosaurus was reinterpreted with a horizontal body posture and fully upright limbs. As an early member of the sauropodomorph lineage it was often depicted as a transitional form between bipedal ancestral dinosaurs and the later quadrupedal sauropods, thought to primarily walk on all fours but also able to run on its hind legs like a hadrosaur – although some studies instead concluded it was fully quadrupedal with a downwards-curling tail that made bipedal movement impossible.

The large numbers of skeletons found together were considered to represent evidence for herding behavior, with groups of Plateosaurus being caught in catastrophic mudflows all at once.

2020s

Extensive biomechanical studies in the 2000s and early 2010s clarified what sort of posture Plateosaurus was really capable of. It was found to be completely unable to position its arms in a quadrupedal stance, and so was actually purely bipedal – and skeletons that had been mounted in the quadrupedal position had needed many of their joints to be completely dislocated to achieve the pose!

A huge number of different Plateosaurus species had been named over the genus’ nearly-200-years of history, too, creating a confusing mess of dubious and invalid names. These were all finally revised in 2019 leaving just three valid species, with Plateosaurus trossingensis as the best known and the new type species.

We now know Plateosaurus lived across central and northern Europe during the Late Triassic, around 214-204 million years ago, at a time when the region had a subtropical climate. It had a small head on a long flexible neck, with teeth convergently resembling those of modern iguanas suggesting it was probably primarily herbivorous (with possible opportunistic omnivory). Its arms were proportionally short for a prosauropod but were well-adapted for grasping, with large claws that may have been used to dig up roots and tear down branches

It had a rapid growth rate and bird-like lungs and air sacs that suggest it was warm-blooded, and different individuals showed an unusually high amount of variation in adult size and age of maturity. Some appear to have been fully grown at about 5m long (~16′) and as young as 12 years old, while others reached 10m long (~33′), and were still growing at 27 years old.

The bonebeds are no longer thought to represent mass mortalities of herds, but instead were probably a scenario more similar to the La Brea Tar Pits – mud-miring traps that smaller lighter animals could escape from but larger individuals became stuck and died.

No prosauropod skin impressions have been found yet, so it’s still unknown whether Plateosaurus was scaly like later sauropods or if it had some degree of protofeather hair-like fuzz.

Shri

About 72 million years ago, in the Late Cretaceous of what is now Mongolia, a dead dromaeosaurid dinosaur lost its head.

30 years ago, in 1991, its headless fossilized remains were discovered during a joint Mongolian Academy of Sciences / American Museum of Natural History expedition in the Gobi Desert.

For a long time the specimen was known only by the nickname of “Ichabodcraniosaurus”, in reference to a character haunted by a headless ghost in the story The Legend of Sleepy Hollow – but now it’s finally been given a full scientific description and a proper name.

Say hello to the first new non-avian dinosaur of 2021, Shri devi!

Named after a buddhist deity, this little dinosaur was around 2m long (6’6″), roughly the size of a modern peacock or wild turkey. It was a very close relative of Velociraptor, but lived in a slightly different part of the ancient Gobi than its famous cousin, giving us a glimpse of how dromaeosaurid species varied across that region.

A map of the Gobi region of Mongolia and China, showing locations where various dromaeosaurid dinosaur fossils have been found. — [ From fig 28 in Turner, A. H. *et al* (2021). A New Dromaeosaurid from the Late Cretaceous Khulsan Locality of Mongolia. *American Museum Novitates*. https://doi.org/10.1206/3965.1 ]

Weird Heads Month #12: Double-Crested Dinosaurs

Dilophosaurus wetherilli is a fairly recognizable dinosaur thanks to its memorable appearance in the Jurassic Park franchise – but unfortunately that also means the popular image of it is completely wrong.

Rather than a small frill-necked venom-spitting creature, this early theropod was actually rather large, reaching around 7m long (~23′), and along with its distinctive double crests it also had a narrow snout with large teeth and a distinctive notch at the front of its lower jaw.

It lived in North America during the early Jurassic, about 196-183 million years ago, and while it wasn’t venomous its notched jaws were probably capable of delivering powerful bites to small struggling prey, much like the similar-looking ornithosuchids in the Triassic. Some structural similarities to the skulls of spinosaurids suggest it may have primarily eaten fish.

Its two bony crests were probably used for visual display, with juveniles only having small crests that fully developed as they matured. They also may have had a more extensive keratinous covering, so it’s not clear what their actual shape and full extent was in life.

Atlasaurus

Sauropod dinosaurs were just generally weird animals, but there’s something… not quite right about Atlasaurus imelakei.

Named after the Atlas Mountains of Morocco where its fossil remains were discovered, Atlasaurus lived during the mid-Jurassic period, around 168-165 million years ago. While it wasn’t the strangest–looking sauropod by any means, compared to other species its body proportions still show a particularly bizarre combination of features, with a slightly bigger head, unusually short neck, and very long slender legs that made up nearly half of its 9m height (29’6″).

It’s sort of the uncanny valley of sauropods. Everything about it is just a tiny bit wrong.

A photograph of an Atlasaurus model. Its been reconstructed very skinny, which only serve to emphasize its weird proportions. — And more shrinkwrapped depictions really don’t help with that. [image source]

Its tall shoulders and sloping back resemble the body plan of brachiosaurids so closely that it was initially thought to be an early member of that group, but more recent studies suggest it may have been part of an earlier evolutionary branch of sauropods known as the turiasaurs – which would mean its brachiosaur-like shape was actually the result of convergent evolution.

But what was it doing with such weird proportions?

…We really don’t know. Other short-necked sauropods seem to have been adapted for feeding on lower vegetation only a couple of meters off the ground, but Atlasaurus’ leggy build would have made it a high browser like the brachiosaurids it was mimicking. Its long legs may also have allowed it to move faster, or given it some advantage navigating over rough terrain, but since no other sauropod ever seemed to evolve this way it must have been doing something particularly unique.

Or perhaps it was just an evolutionary fluke. Maybe part of a lineage that had started adapting to short-necked low browsing, then moved back towards the high browsing niche – and happened to end up lengthening their legs instead of their necks to get the necessary height back.

Kaijutitan

Originating from Japanese monster movies like Godzilla, the word “kaiju” is now often used to refer to giant creatures in general – and so it was only a matter of time before a huge sauropod dinosaur was named after the concept.

Kaijutitan maui* was a titanosaur living in Argentina during the Late Cretaceous, about 89-86 million years ago. It’s only known from fragmentary remains, so its full size is difficult to estimate, but it was probably somewhere in the region of 20m long (66′). Nowhere close to the largest sauropod, but possibly one of the heaviest since it does seem to have been rather chunkily built, with stout limbs and an estimated weight of 40-60 tonnes (44-66 US tons).

* Not named for the Polynesian hero, apparently, but for the initials of the Museo Argentino Urquiza.

Kalligrammatids

Did you know butterflies weren’t the first insects to look like butterflies?

Lepidopterans (the group of insects containing moths and butterflies) have been around since the Late Triassic – but it wasn’t until the diversification of flowering plants during the Cretaceous that recognizable moths would have evolved, and true butterflies didn’t actually appear until the early Cenozoic.

Before then, back in the mid-Jurassic about 165 million years ago, a completely different group of insects convergently evolved remarkably butterfly-like features such as large colorful scaled wings and long sucking proboscises.

Known as the kalligrammatids, these insects were giant members of the lacewing group, related to modern forms like antlions and owlflies. But unlike their predatory relatives the kalligrammatids were specialized pollinators, possibly having a mutualistic relationship with the flower-like cones of bennettitales or the pollination drops of some types of conifers. They seem to have originated in China and were found across Asia and Europe by the Late Jurassic, but a few fossils from South America suggest they were even more widespread and may just have a poor fossil record.

They reached wingspans of up to 16cm (~6″), comparable to some of the largest modern butterflies, and often sported conspicuous anti-predator markings on their wings such as stripes and eyespots – so it’s not surprising that they’re often nicknamed the “butterflies of the Jurassic”.

Rather ironically, the extinction of the kalligrammatids was probably linked to the rise of the flowering plants that the true butterflies would later be so dependent on. As flowers diversified and plants like the bennettitales declined, the kalligrammatids dwindled and disappeared, with the last known fossil record coming from the mid-Cretaceous of Brazil about 113 million years ago.

But while they were around, I do wonder if they also exhibited some similar behaviors – such as mud-puddling for extra nutrients, and specifically the habit of drinking the tears of larger animals that we see in some species. Perhaps some non-avian dinosaurs like this Dilong occasionally put up with kalligrammatids sitting on their faces!

Vespersaurus

For around 50 years some very unusual dinosaur tracks have been found in ancient desert sediments in South America: strange footprints showing the impression of only a single toe, a walking style never before seen in any reptiles.

And recently a fossil of what might be the track maker has actually been found.

Named Vespersaurus paranaensis, this new species lived during the Late Cretaceous of Brazil (~90 mya) and was a member of the noasaurid family of theropods, closely related to the weird-jawed Masiakasaurus from Madagascar.

Measuring about 1.5m long (~5′), Vespersaurus was fairly lightly built with legs proportioned for running – and its feet were absolutely unique. Although it had the standard three main toes of a theropod, it bore its weight entirely on the middle toe and held the other digits off the ground. The two raised toes on each foot also had large knife-like claws which may have been used during hunting, vaguely similar to the sickle claws on the feet of dromaeosaurs. But unlike dromaeosaurs these claws weren’t highly curved or pointed, suggesting Vespersaurus used more of a scratching and slashing technique rather than the raptors’ puncture-and-restraint strategy.

Much like ancient horses, it may have developed its single-toed stance as an adaptation for more efficient fast running, possibly to avoid larger predators or to chase down small fast-moving prey like hopping desert mammals.

The known one-toed fossil footprints are actually slightly older than the Vespersaurus fossil, and similar tracks in Argentina have been found dating back to the Late Jurassic (~150mya), so there may have been a long lineage of “one-toed” desert-dwelling noasaurids in South America that haven’t been found yet.

Glowing Dinosaurs

Many modern birds are capable of seeing into the ultraviolet regions of the electromagnetic spectrum, and some of their non-avian dinosaur ancestors might have had the same sort of vision. And much like their living relatives, that means various parts of their bodies and plumage may also have been UV-reflective and UV-fluorescent.

So here’s a Velociraptor with some speculative UV coloration – although this is just what it would look like to human eyes under a blacklight. What it would actually look like to a creature that can see extra colors is impossible to depict on a screen designed for trichromatic vision!