synapsid – Nix Illustration

Tetracynodon

Therocephalians were a group of synapsids very closely related to – or possibly even ancestral to – the lineage leading to modern mammals. They were a diverse and successful group of carnivores during the latter half of the Permian, but suffered massively during the “Great Dying” mass extinction, with only a handful of representatives making it a few million years into the Triassic.

Tetracynodon darti was one of these rare Triassic therocephalian survivors, living in what is now South Africa around 251 million years ago. Only about 25cm long (~10″), it had slender limbs and strong claws that suggest it was a scratch-digger. Its long snout was lined with pointed teeth, and it was probably an active predator hunting by snapping its jaws at fast-moving prey like insects and smaller vertebrates.

Its combination of small size, burrow-digging habits, and unspecialized diet may be the reason it scraped through the Great Dying when most of its relatives didn’t – but unfortunately it seems to have been a “dead clade walking”, disappearing only a short way into early Triassic deposits.

Pachydectes

Modern mammals are the only living representatives of the synapsids, but back during the Permian there were numerous other evolutionary branches – first the pelycosaurs, and later their descendant the therapsids.

Some of the first non-mammalian therapsids were the biarmosuchians, mid-sized carnivores with a more upright posture than their pelycosaur ancestors. They had large canine teeth in their jaws and powerful bites, and some of them also developed elaborate ornamentation on their skulls, with various bony bumps and crests adorning their faces.

Pachydectes elsi was a 1.5m long (~5′) biarmosuchian living in what is now South Africa during the late Permian, about 265 million years ago. Bone texture indicates its head ornamentation was covered by either tough thickened skin or a keratinous sheath, and the large bulbous bosses on the sides of its snout had a particularly rich blood supply, suggesting these structures could have been continuously growing throughout its entire life.

But despite how well-protected it looked, Pachydectes’ skull was actually relatively fragile and wouldn’t have been able to withstand the impact forces of using its headgear for fighting or defense. Instead it may have been mostly used for visual display – and the blood supply to the snout bosses might even have given it the ability to “blush” them if they had a soft-tissue covering.

Crystal Palace Field Trip Part 1: Walking With Victorian Monsters

This past week I’ve been out of town and unable to work on much art, but instead here’s something a little different. I finally got the chance to go visit some familiar old faces out in the wilds of south London, so let’s go on a little tour of these iconic Victorian-era retrosaurs…

A photograph of an informational sign in London's Crystal Palace Park. The text on it reads, "The Crystal Palace Dinosaurs 1854, a journey through time and science". Three of the iconic Victorian dinosaur statures are also pictured below the title, showing the Iguanodon, Hylaeosaurus, and Megalosaurus.

The Crystal Palace Dinosaurs take their name from the original Crystal Palace, a glass-paned exhibition building originally constructed for a World’s Fair in Hyde Park in 1851.

In 1854 the structure was relocated 14km (~9 miles) south to the newly-created Crystal Palace Park, and a collection of over 30 life-sized statues of prehistoric animals were commissioned to accompany the reopening – creating a sort of Victorian dinosaur theme park – sculpted by Benjamin Waterhouse Hawkins with consultation from paleontologist Sir Richard Owen.

The Palace building itself burned down completely in 1936, and today only the ruins of its terraces remain in the northeast of the park grounds.

Two images of Victorian London's Crystal Palace building. On the left an old black-and-white photograph from around 1854 shows the original structure, a grand glass-paned building with ornate terraced gardens in front of it. On the right a more modern photo from 2011 shows what little remains today – just the ruins of the terraces and stairs. — *The Crystal Palace building then and now*
Left image circa 1854 (public domain)
*Right image circa 2011 by Mark Ahsmann (CC BY-SA 3.0)*

Six sphinx statues based on the Great Sphinx of Tanis also survive up among the Palace ruins, flanking some of the terrace staircases. They fell into serious disrepair during the latter half of the 20^th century, but in 2017 they all finally got some much-needed preservation work, repairing them and restoring their original Victorian red paint jobs.

A photograph of one of the surviving sphinx statues in the Crystal Palace ruins, reclining on a plinth beside some stone steps. It's recently renovated with a coat of terracotta red paint to match its original Victorian-era appearance. In the background the huge Arqiva Crystal Palace telecom tower can be seen.

…But let’s get to what we’re really here for. Dinosaurs! (…And assorted other prehistoric beasties!)

Rechnisaurus

Rechnisaurus cristarhynchus here was a member of the dicynodonts, a group of stocky herbivorous beaky-jawed synapsids that were distantly related to modern mammals. Living in what is now eastern India during the Middle Triassic, about 247-242 million years ago, it’s only known from a single fossil skull – but based on the body proportions of better-known close relatives like Kannemeyeria it was probably somewhere around 1.2m long (~4′).

It had a raised bony crest running down the middle of its snout, with deep bowl-like depressions on either side that probably served to make the crest seem visually larger it already was. (They probably didn’t house any weird soft-tissue structures, however, since these type of dicynodonts tended to have very extensive keratinous coverings over their snouts.)

It also had raised bony areas around its parietal eye, and extensive bony flanges covering most of its tusks giving its face a sort of jowly appearance. All these features were probably for visual display and may have been brightly colored in life.

And, while I usually like to reconstruct dicynodonts as extensively fluffy… recently some fossil specimens of Lystrosaurus have been found showing bumpy leathery skin impressions. This doesn’t necessarily mean that all dicynodonts were hairless (especially since there are still those Permian coprolites), but since kannemeyeriiformes like Rechnisaurus were quite closely related to Lystrosaurus, I’ve gone with no fuzz at all on this one.

Cabarzia

Cabarzia trostheidei here lived during the early Permian in what is now Germany, about 295 million years ago.

Despite its very lizard-like appearance it was actually part of the varanopid lineage, a group of scaly amniotes traditionally classified as early synapsids (distant relatives of modern mammals), but which more recently have been proposed to instead be sauropsid reptiles closer related to early diapsids.

It was around 50cm long (1’8″), and its short arms, long legs, slender body, and long tail suggest it was capable of shifting into a bipedal posture when running at high speeds, similarly to some modern lizards – probably mainly to escape from larger predators, but possibly also used to pursue fast-moving prey like flying insects.

And whether varanopids were actually synapsids or sauropsids, this makes Cabarzia the earliest known example of an animal running on two legs.

Ericiolacerta

The synapsids were an incredibly successful and diverse group during the Permian period, but after the devastating “Great Dying” mass extinction event 252 million years ago only three lineages survived into the Triassic – the cynodonts (close relatives and ancestors of modern mammals), the dicynodonts (beaked tusked weirdos who briefly took over the world), and the therocephalians.

Therocephalians were close relatives of cynodonts, and convergently evolved several very mammal-like anatomical features in their skulls, teeth, and limbs. But unlike their cousins this lineage never fully recovered in the Triassic, and they ultimately disappeared completely around 242 million years ago.

Ericiolacerta parva was one of these short-lived Mesozoic therocephalians, known from the early Triassic (~252-247 million years ago) of South Africa and Antarctica, in regions that were connected at the time as part of the supercontinent of Pangaea. It was a fairly small animal, about 20cm long (~8″), with small sharp teeth that indicate it mainly fed on insects, and semi-opposable thumbs and inner toes that suggest it was also a capable climber.

Holes in the bones of its snout would have carried numerous nerves and blood vessels, which may be evidence of sensitive fleshy lips and possibly whiskers. And while there’s no direct evidence of fur in therocephalians, they do appear to have been active warm-blooded animals – and possible fossilized synapsid hair from the Permian period suggests fuzziness might have been ancestral to all of the “protomammal” lineages that survived into the Triassic.

Retro vs Modern #15: Dimetrodon limbatus

With its prominent sailback Dimetrodon is one of the most iconic prehistoric animals – and one that still frequently gets mistaken for a dinosaur, despite being closer related to modern mammals.

1870s-1980s

The first known Dimetrodon fossil was an upper jaw fragment found in Canada in the 1840s, but at the time this specimen was thought to represent a dinosaur. It wasn’t until the late 1870s that species like Dimetrodon limbatus (initially called Clepsydrops limbatus) from the Midwestern and Southern United States were recognized as belonging to a much older and different group of animals given the name “pelycosaurs“.

While some paleontologists did propose pelycosaurs as being ancestral to mammals quite early on, for several decades the prevailing view was actually that they were an ancient branch of rhynchocephalian reptiles closer related to modern tuataras. From the 1910s onwards pelycosaurs were finally linked back to mammals, with their similarities to the therapsids placing them as early members of the synapsid lineage – although all these early mammal-relatives were still considered to be derived from reptiles, and “mammal-like reptile” became a commonly-used term for them.

As a result reconstructions of Dimetrodon during this time period usually depicted a highly reptilian and heavily scaled lizard-like animal, with a sprawling belly-dragging pose, protruding crocodilian-like teeth, and a highly shrink-wrapped sail on its back modeled on those of some modern lizards. Some earlier images also showed a short stumpy tail, since Dimetrodon‘s longer tail proportions weren’t confirmed until the 1920s.

2020s

During the late 20^th century new classification techniques led to the messy concept of “reptiles” being properly redefined as sauropsids, and synapsids being recognized as an entirely separate non-reptilian lineage of amniotes. Along with new studies and discoveries this has resulted in our understanding of Dimetrodon changing a lot in the last few decades, moving away from a heavily reptilian interpretation and instead letting it be its own weird “protomammal” thing.

We now know there were at least a dozen different species of Dimetrodon living during the early-to-mid Permian, about 295-272 million years ago. Most of them are known from North America, but an additional species discovered in Germany suggests this genus ranged further across Pangaea than previously thought.

Dimetrodon limbatus was one of the larger species, about 3m long (10′), and like other members of the genus it had a tall narrow skull with high-set eyes and two distinct types of teeth in its jaws. The structure of its nasal cavities suggest it had a good sense of smell, and like the related synapsid Ophiacodon it may have had a closer to “warm-blooded” metabolism than previously thought.

It would have had a very poor sense of hearing, however, and probably didn’t even have any visible ears on its head. It may have been functionally deaf to air-borne sounds entirely, only able to detect vibrations by pressing its lower jaw to the ground.

No skin impressions are known for Dimetrodon. Scaly reptile-like skin has been found on varanopids, a group traditionally classified as very early synapsids – but some recent studies have suggested they were actually part of the true reptile lineage, so their extensive scaliness probably doesn’t apply to synapsids like Dimetrodon after all. There is some possible evidence of rows of square or rectangular scale-like scutes on the underside of the belly and tail in pelycosaur-grade synapsids, but otherwise the next-closest known synapsid skin comes from the distantly-related therapsid Estemmenosuchus, which seems to have had smooth glandular skin similar to a hairless mammal.

The characteristic back sail, formed by highly elongated neural spines on the vertebrae, is now thought to have been covered in a different pattern of soft tissue than older reconstructions depicted. The texture of the bone along the spines’ length shows that at the base they were deeply embedded in the back musculature, then further up they were covered by skin webbing, but then at the tips they may actually have been unwebbed and free-standing, giving a much spinier profile.

While the sail was traditionally assumed to be used for temperature regulation, more recently this has started to seem less likely. The sail doesn’t seem to have been quite as well-supplied with blood vessels as previously thought, and there’s a lack of direct correlation between sail size and body size in different Dimetrodon species and age classes. Instead this structure may actually have been used for visual communication and display, and could therefore have been quite flashy and brightly-colored.

Fossilized trackways also suggest that Dimetrodon didn’t move with a low lizard-like sprawling gait but instead with something more like a crocodilian “high walk”, with its limbs much closer to upright. It was probably a fairly active terrestrial predator and would have eaten a wide variety of other smaller Permian animals, with its teeth having been found in association with the remains of the amphibians Eryops and Diplocaulus and the freshwater shark Xenacanthus.

Gordodon

Some of the earliest large terrestrial herbivores on Earth were the edaphosaurids – a very early-branching group of synapsids, the evolutionary lineage whose only modern surviving members are mammals. Like their more famous cousin Dimetrodon these animals sported huge elaborate sails on their backs formed from highly elongated vertebral spines, but despite the similarity in appearance they actually seem to have evolved these structures completely independently.

Known from a single partial skeleton discovered in southern New Mexico, USA, the edaphosaurid Gordodon kraineri dates to around the very end of the Carboniferous or the very earliest Permian, about 299 million years ago.

It was fairly small for an edaphosaurid at about 1.5m long (~5′), and seems to have had transitional anatomy between earlier and later members of the group. Its sail spines were thicker than those of earlier species but still less heavyset than those of later forms, and while each spine had numerous side projections these structures were small, thorn-like, and randomly distributed, unlike the more organized thick crossbars seen in Edaphosaurus.

Its head was proportionally small compared to its body, but still relatively large for an edaphosaurid, and it had an unusually long neck for an early synapsid. But its most distinctive features were its jaws and teeth – it had a narrow snout with a pair of large incisor-like teeth at the front of both its upper and lower jaws, followed by a large toothless gap (a diastema) and then a short row of small peg-like teeth. Like Edaphosaurus it also would have had batteries of grinding tooth plates inside its upper and lower jaws, but probably not as extensively.

Overall its tooth arrangment looked more like a modern herbivorous mammal than an early synapsid, much more highly specialized than anything else known to be alive at the time – the next synapsid known to convergently evolve similar teeth lived around 90 million years later!

It probably had a very different diet to its relatives, with its specialized teeth and fairly slender body suggesting it may have been a selective feeder, cropping the softer more nutritious parts of plants like the fleshy seeds and cones of gymnosperm plants.

Its discovery also hints that herbivorous edaphosaurids in general were much more diverse than we previously thought, and there may be even more surprising forms out there still to be discovered.

Perplexisaurus

If there’s any equivalent to carcinization in mammals, it’s turning into an otter-beaver-like semi-aquatic form.

Because it just keeps happening.

Modern examples alone include otters, beavers, muskrats, giant otter shrews, desmans, aquatic genets, yapoks, lutrine opossums, and platypuses – and in the fossil record there were early pinnipeds, remingtonocetids, pantolestids, stagodontids, and Liaoconodon going as far back as the early Cretaceous. Even outside of the true mammals there were also Castorocauda, Haldanodon, and Kayentatherium during the Jurassic, and much further back in the late Permian there was the early cynodont Procynosuchus.

So a non-cynodont synapsid doing the exact same thing really isn’t all that surprising.

Perplexisaurus foveatus was a member of the therocephalians, a group of synapsids that were close evolutionary “cousins” of the cynodonts-and-true-mammals lineage. Similar in size to a modern rat, about 20cm long (8″), it lived in Western Russia during the Late Permian about 268-265 million years ago.

At the time this region was a river plain with a tropical climate, experiencing seasonal floods that turned the whole area into what’s known as “viesses” (a name based on the abbreviation “V.S.S.” standing for “very shallow sea”), vast shallow lake-seas that persisted for weeks or months at a time.

So this little animal has been interpreted as being semi-aquatic, swimming around and feeding on aquatic invertebrates and tiny fish and amphibians. Its skull had numerous pits around the front of its face, suggesting that it had a highly sensitive snout – probably whiskery, allowing it to hunt entirely by touch in dark murky water, but it’s also been proposed to have possibly had an electroreceptive sense similar to modern platypuses.

Smilesaurus

Despite having a genus name that sounds more like it should belong to a cartoon dinosaur mascot for dental hygiene, Smilesaurus ferox was actually a real gorgonopsian, a predatory synapsid distantly related to modern mammals.

Living in South Africa during the Late Permian, around 259-254 million years ago, Smilesaurus was comparable to a medium-sized dog at around 1m long (3’3″). It had some of the longest sabre-like canine teeth of any known gorgonopsian, proportionally comparable to those of sabertoothed cats – and it may have hunted in a similar manner, using powerful grasping limbs to pin down struggling prey and then dispatching it with slashing bites.

…And it also turns out that when you don’t horribly shrink-wrap a gorgonopsian, you end up with something that looks rather like a bear-hippo.

(For some similarly chonky gorgonopsians, check out Tas’ @i-draws-dinosaurs reconstructions here. Bullet Man was definitely a bit of an inspiration in this.)