Weird Heads Month #10: Permian Crowns

The tiny-headed Cotylorhynchus we saw earlier in this series wasn’t the only synapsid with a weird head.

A little more closely related to modern mammals, the dinocephalians were a a diverse group that were found across Pangaea during the middle of the Permian period. Many of them had thickened skulls that may have been used for headbutting each other, and some also developed bony horn-like projections around their faces.

And Estemmenosuchus mirabilis here was particularly elaborately ornamented, earning it a name meaning “wondrous crowned crocodile”. It lived in the Perm region of Russia during the mid Permian, about 268-265 million years ago, and was one of the largest dinocephalians, reaching at least 3m long (9’10”).

It had two big antler-like structures on its head, two wide cheek flanges, and a small nose horn, almost looking like the synapsid version of a ceratopsid dinosaur – and with its big bulky body, fairly erect-legged posture, and herbivorous-or-omnivorous diet it may have been a fairly close ecological equivalent to them, too.

But it’s also possible it was semi-aquatic, and it certainly does have a very hippo-like appearance when reconstructed with a decent amount of soft tissue.

One specimen of Estemmenosuchus even preserved skin impressions around its face, which were described in Russian in the early 1980s. They show scaleless glandular skin with a slightly bumpy texture, similar to that of hairless mammals or some amphibians. Since it occupied a point in the synapsid family tree close to where hair may have originated (somewhere in the Permian therapsids), it’s not clear if it was entirely hairless or if it had just secondarily lost some of it.

Weird Heads Month #02: Tiny Heads

Sometimes the really weird thing about a head isn’t any sort of ridiculous ornamentation.

Sometimes it’s just the wrong size.

That’s what was going on with Cotylorhynchus romeri from the early Permian of North America, living about 280-272 million years ago. Despite looking like a big fat lizard this creature was actually a very early synapsid, closer related to modern mammals than to reptiles, and it was a distant cousin of other stem-mammals like the famous Dimetrodon.

Around 3.5m long (11’6″), it was one of the largest herbivores of the early Permian, with a very wide barrel-shaped body, chunky limbs, and a comically small head. Such a tiny head isn’t necessarily unique – another synapsid Edaphosaurus also had a fairly small skull compared to its body, and dinosaurs like stegosaurs, sauropods, and moa had heads even more disproportional. But something about Cotylorhynchus in particular just looks… incredibly odd.

It also had some surprisingly sizeable nostril openings in that little skull, and it had may have had a very good sense of smell or perhaps some sort of specialized breathing system like the modern saiga’s “air conditioning” nose.

Although usually depicted as a fully terrestrial animal, the structure of Cotylorhynchus‘ bones and its flattened paddle-like hands and feet have recently been used to suggest that it may have been semi-aquatic, more of a Permian hippo than a cow. But such a lifestyle would have required it to have a much more efficient method of breathing than previously thought – suggesting it had a mammal-like diaphragm, and possibly also explaining that weird nose.

Tiarajudens

Synapsids just keep evolving saber teeth.

Both proto-mammals and true mammals have independently evolved oversized fangs quite a few different times in a lot of different lineages over the last few hundred million years (even in some modern ones), and one of the first to experiment with such a feature was Tiarajudens eccentricus.

Living in southern Brazil towards the end of the Permian period, about 265-260 million years ago, Tiarujudens was an early member of a group of known as anomodonts. These chunky herbivorous synapsids weren’t directly ancestral to modern mammals, but were instead evolutionary cousins, and their lineage eventually included tusked dicynodonts like the world-conquering Lystrosaurus.

Tiarajudens was around 1-1.2m long (3’3″-3’11”) and sported a pair of very long blade-like canine teeth in its upper jaw. Since the rest of its teeth were clearly adapted for eating plants – with one of the the earliest known examples of flat grinding molars that would have allowed it to chew up tough vegetation – these fangs probably served more of a display or defensive function.

The saber teeth may even have been a sexually dimorphic feature like in modern musk deer. Another anomodont from South Africa, Anomocephalus africanus, is incredibly similar to Tiarajudens except for a lack of fangs – and since South America and Africa were connected as part of Pangaea at the time, it’s possible that these two actually represent males and females of the same species.

Without finding a larger number of fossils we can’t know for certain, but it’s an interesting possibility at least.

Remigiomontanus

Edaphosaurids were a fairly early branch of the synapsids – the evolutionary lineage whose only surviving members are modern mammals – and were some of the earliest known tetrapods to develop into large specialized herbivores. They also had huge spiny sails on their backs resembling those seen in their cousins the sphenacodontids (including the famous Dimetrodon), but the two groups actually evolved those features completely independently of each other.

Although their fossils are known from both North America and Europe, their European remains are very rare and fragmentary. Currently the best-known specimen is made up of a recently-discovered partial spinal column and a few hand and tail bones.

Given the name Remigiomontanus robustus, this edaphosaurid lived in western Germany during the end of the Carboniferous and the start of the Permian, around 300-298 million years ago. About 1.2m long (3’11”), it seems to represent an intermediate form between small insectivorous-or-omnivorous edaphosaurids like Ianthasaurus and the huge herbivorous Edaphosaurus.

(Interestingly the paper that names Remigiomontanus also makes a brief mention that the protruding cross-bars on edaphosaurid sails may have anchored larger keratinous coverings, which could have made them look even more spectacularly spiky and suggests their sails may have served an anti-predator function. Hopefully if this is true we’ll see further information get officially published about it sometime!)

Diademodon

Modern mammals are the only surviving members of a much larger evolutionary group known as the synapsids – which back in the Permian period were the dominant land vertebrates.

But much like all other life on Earth at the time, the synapsids were absolutely devastated in the “Great Dying” mass extinction at the end of the Permian, 252 million years ago. Only three lineages survived into the Triassic: the dicynodonts (who briefly took over the world), the therocephalians (who went extinct not long afterwards), and the cynodonts (who eventually gave rise to early mammals).

Diademodon tetragonus here lived right in the wake of the extinction during the Early and Middle Triassic, about 251-242 million years ago. Around 2m long (6′6″), it was one of the largest known cynodonts, and it must have been a fairly successful species since it ranged across a large chunk of Pangaea, known from modern southern Africa, South America and Antarctica.

It had pig-like cheekbones and enormous jaw muscles, along with sharp incisors and canine teeth at the front of its jaws and grinding molars at the back. This arrangement suggests that much like modern pigs it may have been an opportunistic omnivore, occasionally snacking on smaller animals and carrion – although an isotope analysis of its teeth indicates the vast majority of its diet was probably still vegetation in shady damp environments.

Lisowicia

Dicynodonts were a group of herbivorous animals with toothless beaks and protruding tusks, part of the synapsid lineage and much closer related to mammals than to reptiles. They were some of the most successful and widespread land vertebrates from the Late Permian to the Middle Triassic, with one genus even briefly taking over the world in the aftermath of the End-Permian mass extinction event.

And it turns out some of them got very big.

Fossils of a surprisingly large dicynodont were first reported in 2008, but it wasn’t until just recently (in late 2018) that this giant creature was finally given an official name – Lisowicia bojani.

Close in size to a modern elephant, at around 2.6m tall (8′6″) and 4.5m long (14′9″), it was by far the largest known example of its kind to have ever lived. And while most other dicynodonts had upright hindlimbs and sprawling forelimbs, Lisowicia seems to have developed a fully upright posture much more similar to that of quadrupedal dinosaurs and modern mammals.

It was also one of the very last of its kind, living during the Late Triassic of Poland, about 208 million years ago (although there was a possible later survivor in Australia). This was around the same time that early sauropod dinosaurs were likewise first starting to experiment with gigantism, suggesting that both groups were convergently evolving to exploit newly-available ecological niches.

Month of Mesozoic Mammals #08: Tiny Transitional

Hadrocodium

Falling evolutionarily between the docodonts and Mammalia itself, Hadrocodium is an important transitional form in the early mammal family tree. Something very similar to it would have been the common ancestor of all modern mammals.

Living in China during the Early Jurassic (196-189 mya), it was one of the first known mammals to have both an enlarged brain cavity and the characteristic middle ear bones of modern mammals, about 45 million years earlier than such traits were previously thought to have evolved.

It was also one of the smallest mammals of all time, measuring only about 3cm long (1.2″) – similar in size to the smallest mammals alive today, the bumblebee bat and Etruscan shrew.

Month of Mesozoic Mammals #07: Digging Down

Docofossor

The docodonts didn’t stop at exploiting ecological niches in the trees and water. Another branch of the group specialized into underground burrowing, developing convergent features remarkably similar to modern golden moles.

Docofossor is known from the Middle Jurassic of China (161-155 mya), and measured about 10cm long (4″). It had large shovel-like fingers, strong forelimbs, short sprawling hindlimbs, and pointed teeth adapted for capturing invertebrate prey. (I’ve also given it a patch of protective keratinized skin on its snout here, based on the related Haldanodon.)

It had a reduced number of bones in its fingers, a modification identical to some modern mammals – suggesting that these relatively “primitive” mammals were already using the exact same genes to regulate their anatomical development.

Month of Mesozoic Mammals #06: Going for a Swim

Castorocauda

While some docodonts like Agilodocodon were going up into the trees, another branch of the group was specializing into semi-aquatic habits instead.

Castorocauda is known from the Middle Jurassic of China (165-161 mya), represented by an exceptionally preserved fossil showing soft tissue and hair impressions. About 40cm long (1′4″), it would have lived in a wetland environment and was well-adapted for swimming, with a flattened scaly beaver-like tail, webbed toes, and a coat of dense fur very similar to that of modern mammals, made up of both guard hairs and underfur.

Its strong forelimbs suggest it was capable of digging burrows, like modern platypus, and its sharp backwards-pointing teeth indicate a diet of slippery prey such as fish and worms.

It was also one of the earliest known mammals with (possibly venomous) spurs on its ankles. This feature is only seen today in monotremes, but seems to have been an ancestral trait common to all early mammals that was later lost in the lineage leading to marsupials and placentals.

Month of Mesozoic Mammals #05: Climbing Trees

Agilodocodon

Before we get to the actual-Mammalia-mammals, there’s one more group of mammaliaformes who deserve some attention – the docodonts.

Falling evolutionarily just outside of Mammalia itself, docodonts first appeared in the mid-Jurassic and lasted until the Early Cretaceous. They used to only be known from teeth and jaw fragments and were thought to have been fairly generic shrew-like terrestrial insectivores, but more recent discoveries have shown them to have actually been some of the earliest mammals to specialize into diverse habitats.

Agilodocodon was adapted for climbing around in trees, making it one of the earliest known arboreal mammals (although not the first climbing synapsid). Living in China during the Middle Jurassic (165-161 mya), it measured about 13cm long (5″) and had sharp gripping claws and flexible wrists and ankles similar to modern climbing mammals like tree squirrels.

When it was first described in 2015 it was suggested that its spade-like front teeth were specialized for gnawing bark and feeding on tree sap – but a later study found that its teeth didn’t really resemble those of any modern sap-eating mammals, and in fact were closer in shape to those of insectivorous marsupials and elephant shrews.