thyreophora – Nix Illustration

Strange Symmetries #11: Step Up To The Plate

Stegosaurs are some of the most popular and recognizable dinosaurs thanks to their unique appearances, with small heads, elaborate back plates, and spiky thagomizer tails.

Closely related to the ankylosaurs, they first appeared in the mid-Jurassic about 170 million years ago. While they lasted until at least the mid-Cretaceous (~100 milion years ago), their heyday was in the latter half of the Jurassic, ranging all across Asia, Europe, Africa, and North America – and the North American species like the eponymous Stegosaurus developed especially elaborate plates in a distinctive asymmetrical pattern, not arranged in pairs like most other stegosaurs but in alternating rows along each side of the midline of their backs.

Hesperosaurus mjosi lived around 156 million years ago during the late Jurassic, in what is now Wyoming and Montana in the Western United States. It was closely related to Stegosaurus but was a little older and a little smaller, about 5-6m long (~16-20′).

Much like its more famous relative its plates seem to have alternated along its back, which may have been an adaptation to maximize visible surface area while minimizing the number of plates, saving on the energy needed to grow such large elaborate ornamentation.

Hesperosaurus might also represent a rare case of possible sexual dimorphism in non-avian dinosaurs, with wider more rounded plates potentially interpreted as belonging to males and taller pointed plates belonging to females.

Jakapil

The thyreophorans were heavily armored ornithischian dinosaurs, with their most famous representatives being the stegosaurs and the ankylosaurs. Earlier members of the group were all small bipedal animals covered in rows of prickly osteoderms, and until now these “primitive” forms were known only from the early-to-mid Jurassic, around 200-165 million years ago.

But now the recent discovery of Jakapil kaniukura is suggesting a lineage of early thyreophorans actually survived for much much longer than previously thought – all the way into the Late Cretaceous, about 97-94 million years ago.

Just 1.5m long (5′), Jakapil lived in what is now southern Argentina, in an ancient desert with a braided river system. It was bipedal, with a short beak, small arms, and a body bristling with spiky armor, and its unusually deep lower jaw and heavily worn teeth indicate it fed on rather tough vegetation that required a lot of chewing to process.

It’s currently only known from somewhat fragmentary remains, so reconstructions of its full appearance are rather speculative and there’s already been some dispute about whether Jakapil actually was a thyreophoran. One proposal is that it shared a lot of anatomical features with early ceratopsians instead, which if true would make it an incredibly weird armored ceratopsian, and also the first definitive member of that group from South America. But the ceratopsian-like features could also just be due to convergent evolution – and a Jakapil-like dinosaur might actually help explain the only other known dubious South American “ceratopsian” Notoceratops, and the similarly-disputed Australian Serendipaceratops.

But whatever it was – late-surviving basal thyreophoran, southern armored ceratopsian, or even a previously unknown lineage of ornithishcians entirely new to science – it’s an exciting and unexpected discovery.

Retro vs Modern #13: Stegosaurus stenops

The first known stegosaur fossils were found in England and South Africa between the 1840s and 1870s, but these dinosaurs weren’t properly recognized as a highly distinctive group until the discovery of Stegosaurus itself in North America during the late 1870s.

1880s

The first Stegosaurus reconstructions were based on fragmentary and disarticulated fossil material, and its life appearance was very poorly understood. Initially it was depicted as a bipedal long-necked animal, with its plates laying flat against its back like a turtle shell, numerous spikes across its back, and more plates running along its tail.

1890s-1970s

Better skeletons of the species Stegosaurus stenops were discovered in the late 1880s, and by the 1890s stegosaur anatomy was becoming clearer. Reconstructions quickly adopted an arch-backed body shape with a tiny head and drooping tail, short semi-sprawling forelimbs and long hindlimbs, and with the plates now properly upright on the back and the spikes at the end of the tail.

Stegosaurus‘ unique appearance rapidly made it one of the most famous and recognizable dinosaurs to the general public. Its comical-seeming tiny head and even tinier brain also unfortunately ended up contributing to the prevailing early 20^th century attitude that dinosaurs were sluggish and unintelligent, with the myth that it needed a “second brain” in its hips to control its huge body becoming a popular notion for quite some time.

The exact arrangement of the iconic back plates and tail spikes was uncertain for several decades, with early versions in the 1890s having up to eight tail spikes and a single row of plates. This was then updated in the 1900s to a double row of symmetrical plate pairs, and by the 1920s the standard arrangment had soon become an alternating two-row pattern with the tail spikes reduced to four – a layout that’s still considered correct today.

2020s

In the second half of the 20^th century a combination of numerous new stegosaur species from China and the Dinosaur Renaissance began to revise the way Stegosaurus was understood, bringing it into a fully upright posture with its head and tail held high, and recognizing the convergently sauropod-like anatomy of its hands and feet.

But something still wasn’t right.

Compared to other known stegosaurs, Stegosaurus itself was starting to seem… rather weird. Its short neck, short forelegs, giant plates, sloping back and high rump were much more exaggeratedly proportioned than any of its relatives.

This was finally resolved in the 2010s when a near-complete specimen nicknamed “Sophie” was thoroughly described – and revealed that Stegosaurus’ proportions had been wrong the whole time. All previous skeletal reconstructions had been composites, put together from remains of multiple individuals that had all been different ages and sizes, and in the process had heavily distorted our idea of what this animal actually looked like.

Our modern view of Stegosaurus is now a much more typical stegosaur than before. It lived during the Late Jurassic, about 155-145 million years ago, across the Western and South Central United States (with a possible additional occurence in Portugal), alongside several other iconic dinosaurs of the “Jurassic savanna” like Brontosaurus, Diplodocus, and Allosaurus.

It grew up to around 9m long (~30ft), and had a small head with a long narrow snout, with a toothless beak at the front of its jaws and small peg-shaped teeth further back. Bony ossicles lined the underside of its neck, possibly providing chainmail-like protection to its throat, and its skin was covered in tiny pebbly scales interspersed with “rosettes” formed around slightly larger oval scales. Its neck was longer than previously thought, more in line with other stegosaurs, and its torso and hind legs were a bit shorter, making its posture more horizontal and its back less arched.

The actual function of the large back plates is still uncertain. Ideas about them being defensive armor (and speculation about them even being moveable!) have mostly been discounted at this point, since they were actually relatively fragile – although their keratinous covering may have had a fairly sharp edge. Thermoregulation has been a popular explanation for many decades, with blood vessel impressions in the plates being proposed as evidence they were used as “radiators” to prevent overheating like the ears of modern African elephants.

But currently the most likely primary plate function is thought to be visual display, with the large plates increasing the perceived size of Stegosaurus either to intimidate predators and rivals or to impress potential mates. If this was the case then they may have also been strikingly colored and patterned in life.

Meanwhile the “thagomizer” on its tail actually does seem to have been a weapon, with injuries to that area of the body being fairly common, and several Allosaurus fossils have been found with puncture wounds the exact size and shape of Stegosaurus spikes. Articulated specimens have also shown that the tail curved downwards at the tip, holding the thagomizer with the spikes pointing horizontally outwards and backwards.

Retro vs Modern #03: Hylaeosaurus armatus

Despite being the third-ever scientifically named dinosaur genus, and being used in the first official definition of dinosaurs as a group, Hylaeosaurus armatus has ended up as a much less well-known name than Iguanodon or even Megalosaurus.

It was also the very first ankylosaur to be discovered, found as a partial skeleton in Southeast England in the early 1830s. Its large bony spikes were quickly recognized as being some sort of defensive armor, initially thought to be arranged in a vertical row along the animal’s back and tail.

1850s

The Victorian Crystal Palace statue of Hylaeosaurus is surprisingly decent for such an early attempt at reconstructing something as weird as an ankylosaur. It gives the impression of a slower and much more lizard-like animal than Iguanodon or Megalosaurus, showing it as a large squat quadruped with hoof-like claws and heavily armored scaly skin, with long spines running along its back and numerous smaller bony bumps over its head and sides.

1860s-1920s

Discoveries of other more complete armored dinosaurs began to give a better picture of what ankylosaurs actually looked like. But although Hylaeosaurus was soon recognized as having had multiple rows of spikes rather than just one, actual reconstructions of it seem to have been scarce during this period – mostly all derivative of a single 1869 image that depicted it as a sort of fat sprawling pinecone-lizard bristling with spikes.

2020s

Still only known from fragmentary material, Hylaeosaurus has remained rather obscure for a long time. In the 21^st century it’s started to get a bit more attention, however, with the original specimen being further prepared and examined – and 2020 was Hylaeosaurus’ big year, with both a redescription of the genus being published and it also being featured on a special-edition British 50p coin.

Hylaeosaurus was probably around 4m long (~13′), and lived in southeast England about 140-136 million years ago. It may have also ranged further across Europe, with possible remains known from Germany and some more dubious records from France, Spain, and Romania. Generally classified as an early nodosaurid, most of our modern knowledge of what it would have looked like comes from other discoveries of much better-known ankylosaurian relatives, including some exquisitely well-preserved examples in the last few years like Borealopelta and Zuul.

It would have had a low triangular head, with a toothless beak at the front of its jaws and leaf-shaped teeth further back, and a pair of short horns on the back of its skull behind its eyes. Rows of spiky osteoderm armor ran along is body, with longer curving spines over its shoulders, all covered in thick keratin sheaths that would have made them look even larger in life. Numerous smaller bony nodules in its skin filled in the gaps between the armor, forming a tough protective shield over its entire back. Its short powerful limbs had hoof-like claws, and if it was indeed a nodosaurid its tail would have lacked the famous club of its ankylosaurid cousins.

Based on Borealopelta we even now know a little bit about the potential coloration and patterning of these animals – some of them were reddish-colored, with a countershaded camouflage pattern, darker on top and lighter on the underside.

Stegouros

While some ankylosaurs are famous for their specialized tail clubs, Stegouros elengassen here had something else entirely going on with its rear end.

Known from the late Cretaceous of southern Chile, about 75-72 million years ago, this small ankylosaur was around 1.5m long (~5′), roughly the size of a large dog. It had a proportionally larger head and more slender limbs than most other ankylosaurs, and a pelvis more resembling a stegosaur, but its most distinctive feature was its tail – it had a completely unique never-before-seen type of tail weapon, with a flat “frond-like” structure formed from several pairs of large fused osteoderms making a shape resembling a macuahuitl.

It seems to have been part of a previously unrecognized very early-branching lineage of Gondwanan ankylosaurs – the parankylosaurians – with its closest relatives Antarctopelta and Kunbarrasaurus also included in this new group. And since the tail regions of both of those other species are poorly known, this means they may also have possessed macuahuitls.

Eons Roundup 9

New year, new PBS Eons commission roundup day!

An illustration of an extinct early walrus. Unlike modern walruses it has no tusks, more resembling a sea lion in appearance. — Neotherium mirum

An illustration of an extinct species of walrus. It has shorter thicker tusks than modern walruses, and it's depicted grubbing around on a muddy seafloor for buried shellfish. — Neotherium mirum

The ancient walruses Neotherium and Valenictus, from “How the Walrus Got Its Tusks”
https://www.youtube.com/watch?v=BKDGYGV2LK8

An illustration of an extinct nodosaur, a type of quadrupedal herbivorous dinosaur. Its body is covered in densely-packed knobbly spiky armor. — Borealopelta markmitchelli (life restoration)

An illustration of the carcass of an extinct nodosaur, a type of quadrupedal herbivorous dinosaur. Its body is covered in knobbly spiky armor, and its belly is massively bloated up from trapped gasses, forcing it to have turned upside down. — Borealopelta markmitchelli (life restoration)

The nodosaurid ankylosaur Borealopelta, in both alive and “bloat-and-float” carcass states, from “The Dinosaur Who Was Buried at Sea”
https://www.youtube.com/watch?v=a-UZXBF63z4

An illustration of an extinct ankylosaur, a type of quadrupedal herbivorous dinosaur. Its body is covered in knobbly spiky armor, but unlike most other ankylosaurids it has no bony club at the tip of its tail. — Gobisaurus domoculus

An illustration of an extinct ankylosaur, a type of quadrupedal herbivorous dinosaur. Its body is covered in knobbly spiky armor, and it has a small bony club at the tip of its tail. — Gobisaurus domoculus

The ankylosaurid ankylosaurs Gobisaurus and Dyoplosaurus, from “How Ankylosaurs Got Their Clubs”
https://www.youtube.com/watch?v=lRt-4SdzWrk

Scutellosaurus

The distinctive armored ankylosaurs and stegosaurs were very closely related to each other, and were part of a group of dinosaurs known as the thyreophorans.

One of the earliest known members of this lineage was Scutellosaurus lawleri. Living in Arizona during the Early Jurassic, about 196-183 million years ago, it was a small lightly-built bipedal herbivore, only about 1.2m long (3′11″) – with over half that length being just its unusually long tail.

Its body was covered in rows of hundreds of small bony osteoderms, helping to protect it against larger predators like Dilophosaurus. And this was obviously an evolutionary strategy that worked very well for Scutellosaurus and other early thyreophorans, because within about 20 million years they’d given rise to the first true ankylosaurs and stegosaurs – with the tank-like ankylosaurs being especially successful, spreading to every continent and lasting all the way up until the end-Cretaceous mass extinction.

Zuul

Zuul crurivastator, an ankylosaur from the Late Cretaceous of Montana, USA (~75 mya).

One of the most complete ankylosaurids ever found in North America, it’s known from a full skeleton about 6m long (20′). Much of its bony osteoderm armor is preserved in life position, along with skin impressions and the remains of keratinous scales and spike sheaths – although so far only the skull and tail have actually been fully prepared and described.

(The fuzz on this reconstruction is highly speculative, but since it’ll likely end up inaccurate anyway once of the rest of the body is fully described… why not have some fun with it?)

Its genus name was inspired by its skull’s resemblance to Zuul the Gatekeeper from the 1984 movie Ghostbusters, while its species name translates to “destroyer of shins” in reference to its especially large tail club.

Unsolved Paleo Mysteries Month #16 – Strange Snoots 2: Oddball Ornithischians

Those extinct horses weren’t the only ancient creatures with unexplained noses. Some dinosaurs had equally weird things going on with their snouts – and while hadrosaurs’ big honkin’ snoots are fairly well-known, there were other ornithischians with their own bizarre nasal anatomy.

An illustration of the skull of an extinct horned dinosaur, showing the unusually large nasal cavity. Below is a reconstruction of the dinosaur's head in life. — *Triceratops horridus* skull and head reconstruction

Many ceratopsids had an enormous nasal opening forming a giant bony “window” through their snout, with the chasmosaurines like the famous Triceratops having additional bony projections and hollowed regions within these holes. They probably supported some huge elaborate cartilage structures in life, but what they were for is still a mystery. They may have helped with heat dissipation or moisture conservation, aided sound production, provided a highly sensitive sense of smell, housed a vomeronasal organ, held part of an air-filled pneumatic system… or, getting more speculative, possibly even some sort of inflatable nasal display structure.

An illustration of the skull of an extinct armored dinosaur, showing the multiple holes inside the nasal cavity. Below is a reconstruction of the dinosaur's head in life. — *Minotaurasaurus ramachandrani* skull and head reconstruction

Some ankylosaurids, meanwhile, went with multiple holes instead. Minotaurasaurus here had two additional openings around its nostrils, and Pinacosaurus could have up to five – the purpose of which is unknown. Many ankylosaurs also had forward-facing nostrils (a rare trait in archosaurs) and incredibly complex looping airways through their skulls. These may have allowed for mammal-like “air conditioning”, regulating the heat and moisture content of each breath, or perhaps enhanced their sense of smell or served some sort of resonance chamber function. Or, again, maybe even nose balloons.

~~Also floofy ankylosaur because I can.~~