Haldanodon

The docodonts were a group of mammaliaformes (close relatives of the earliest true mammals) which lived across North America, Europe, and Asia from the Middle Jurassic to the Early Cretaceous. Originally only known from teeth and jaw fragments they were traditionally thought to be fairly generic shrew-like insectivores, but more recent discoveries of better fossils have revealed they were actually much more diverse, occupying ecological niches ranging from squirrel-like tree-climbers to mole-like diggers to beaver-otter-like swimmers.

Most of the more complete fossil material of these animals comes from the mid-Jurassic of China, but one species from elsewhere is also known from a partial skeleton.

Haldanodon exspectatus here lived in central Portugal during the Late Jurassic, about 155 million years ago. Around 15-20cm long (6-8″), it had small eyes and short chunky well-muscled limbs with the front paws adapted for digging. Since it inhabited a very swampy environment it probably wasn’t a pure mole-like burrower – extensive tunnels would have constantly flooded – but it may have instead been a similar sort of semi-aquatic animal to modern platypuses and desmans, foraging for invertebrates in the water and excavating burrows in the banks.

Roughened areas of bone on its snout may also have supported a patch of tough keratinous skin, which would have helped protect its face while digging.

Diictodon

Even before Lystrosaurus briefly took over the world, the dicynodont synapsids were a highly successful group. These herbivorous beaked-and-tusked mammal-cousins were abundant all over the supercontinent of Pangaea during the Late Permian and occupied a wide variety of ecological niches, ranging from rat-sized to almost bear-sized. (And later during the Triassic some got even bigger.)

Towards the smaller end of that size range were species like Diictodon. Living around 259-254 million years ago in Southern Africa (but with fossils also found in northern China, suggesting a much larger geographic range) this dicynodont grew up to about 45cm long (1’6″) and was a gopher-like creature adapted for digging, with a tubular body and short muscular limbs.

It was a very common animal, making up around half of all vertebrate fossils in some locations. Numerous preserved spiral-shaped burrows have been found concentrated in small areas, going down as much as 1.5m (5′) into the ground.

Several different species have been named within the Diictodon genus, but currently they’ve all been lumped together under the single name of Diictodon feliceps. There’s a lot of anatomical variation between specimens, though, with some notably being smaller and lacking the distinctive tusks seen in others – which may be evidence of sexual dimorphism, with the tuskless individuals possibly being females. (Although differences in inner ear anatomy may also indicate they were a separate species entirely, in which case female D. feliceps might instead be represented by fossils showing smaller tusks.)

I’ve illustrated one of the tuskless forms here, since they don’t generally get as much attention as the tusked ones. It’s also speculatively fluffy and iridescent similar to modern golden moles.

Weird Heads Month #30: Lumpy-Faced Synapsids

Among the synapsids (“proto-mammals”), head ornamentation evolved multiple times in the therapsids, from basal members of the group like Tetraceratops, burnetiamorphs, and dinocephalians to later lineages like dicynodonts and gorgonopsids.

But these sorts of structures don’t seem to have really ever developed in one of the lineages most closely related to the ancestors of modern mammals, a group known as therocephalians.

…With the exception of Choerosaurus dejageri.

Living in South Africa during the late Permian, around 259-254 million years ago, this small synapsid was only about 35cm long (1’2″) but sported some large bulging bony bosses on the sides of both its snout and lower jaw.

The bosses would have been covered by tough skin in life, similar to modern giraffid ossicones.

A study of Choerosaurus‘ skull found that its head was rather delicately built, and the bosses were relatively fragile and lacked the sort of reinforcement needed to resist impacts, suggesting that these structures weren’t used as weapons for fighting each other but were probably more for display – so they may even have been brightly colored.

The upper jaw bosses were also well-supplied with nerves and blood vessels, and would have been quite sensitive to touch.

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.

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.

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.

Procynosuchus

Procynosuchus delaharpeae, a synapsid from the Late Permian (~259-252 mya). Measuring about 60cm long (2′), it was one of the earliest members of the cynodonts, the lineage that would eventually lead to mammals.

Its fossils are mostly known from southern Africa, but similar remains have also been found in Europe and Russia, suggesting it was actually quite widespread across the supercontinent of Pangaea that existed at the time.

It had a long vertically-flattened tail, strong leg muscles, and paddle-like feet – all adaptations that suggest it was a semi-aquatic otter-like animal capable of agile swimming. It also had forward-facing eyes, giving it good binocular vision and depth perception while pursuing fish underwater.

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.