You won't believe how interesting the Mesozoic mammaliaform Morganucodon watsoni actually is!

Morganucodon watsoni forage for insects and spiders at night among Carboniferous limestones in early Mesozoic Wales. Empty, torn webs suggest they're having a good night, while forest fires burn in the background. Prints are available.

Vertebrate palaeontology textbooks always devote a good chunk of space to morganucodonts, a clade of Triassic-Jurassic Mammaliaformes representing some of the first wholly 'mammal-like' synapsids. Even folks largely uninterested in Mesozoic synapsids will be familiar with two morganucodonts - the Triassic, South African Megazostrodon and the widely-spread Triassic-Jurassic Laurasian genus Morganucodon - because they have become the quintessential 'Mesozoic mammal': small, shrew-like creatures which scurried around the feet of reptiles. Their lack of unusual proportions or adaptations for specialist lifestyles has probably played a part in Mesozoic mammaliaforms being considered a bit boring compared to their dinosaur contemporaries (sorry, mammal palaeontologists), a feat not helped by a deficit of new, particularly interesting artwork of them. Morganucodont representations in palaeoart are frequently quite 'diagrammatic' with 'generic' early Mesozoic backgrounds, animals in lateral or anterolateral aspect to show off their anatomy and daylight settings, despite frequent discussion about their possible/likely nocturnal habits.

Recently, I was asked to produce a reconstruction of Morganucodon watsoni for some friends (above). Being better versed in reptiles than mammals, I had to undertake a fair bit of research to bring myself up to speed on morganucodont anatomy and evolution. Turns out that reputation for being dull is entirely unwarranted: Morganucodon, its relatives, and the world they inhabited are really fascinating. Indeed, they should be bona fide fossil celebrities: Morganucodon and Megazostrodon were the first Mesozoic mammaliaforms known from anything like complete remains (albeit lots of fragments for the former, rather than a single complete specimen) instead of mere teeth and jaws. Their discovery in the mid-20th century can be seen as the start of a new era of understanding of Mesozoic mammaliaform life, and a forerunner of our now rapidly expanding knowledge of Mesozoic mammal diversity.

Initially, I wanted to write a long, detailed post about this painting and the animals it depicts, but that just won't be happening this side of 2015. But, dammit, Morganucodon is too cool to ignore completely or short-change with a 'picture of the day' post, so here's a quick-fire, from-the-hip summary of the research behind the work.

We'll start with the setting. Fossils of M. watsoni occur in British Triassic/lower Jurassic 'fissure fills' cropping out around Bristol and south Wales. These deposits represent ancient infills of caves carved into much older Carboniferous limestones, which are perhaps best known for yielding the sauropodomorph Thecodontosaurus. Fissure fill outcrops occur in multiple quarries across the southern UK and are frequently rich in fossil material, but the mammaliaforms are rare components of the fauna constrained to just a few localities. Dating the fissure fills is not easy because they contain few fossils useful for dating. Some quarries are reliably set at the Rhaetian, but they are probably not all of the same age: some may be as young as the earliest Jurassic. These include sites which contain M. watsoni fossils, which is why different texts give slightly different ages for this animal.

The upper Triassic/lower Jurassic of Europe would be an ideal holiday location for many. The desert landmass of Pangaea was in the process of breaking up, and Europe - including the southern UK - was in the process of being flooded by shallow seas. By the time M. watsoni appeared, Britain's only landmasses were small, low-lying, forested islands which, in terms of climate and general topography, would have resembled those of the Caribbean. Burned plant remains in the fissure fills indicate that the forested inlands of these islands burned on occasion, the remnants of forest fires being washed into coastal limestone caves by storms and floods. We can identify the fissure fill caves as coastal because they contain marine fossils along with terrestrial and freshwater species. It's on these limestones that I set this painting, a deliberate move to avoid another cycad-filled 'semi-arid' Triassic scene.

The island home of M. watsoni was ruled by reptiles - but not necessarily the ones you expect. Although dinosaurs were common, represented by both sauropodomorphs and theropods, the most abundant and diverse reptile group were sphenodonts. These guys deserve their own posts and paintings one day, their Mesozoic run being far more than just spreading tuatara clones across the world. Anyway, the decaying body of one of these - Planocephalosaurus - can be seen in the foreground of the painting above. As these indicate, the sphenodonts contemporary with M. watsoni weren't huge, but they still likely gave the resident mammaliaforms a hard time in competing for similar food resources. It's interesting to ponder how these animals carved up their respective ecologies to avoid direct competition with one another.

Scientists predict that, if Morganucodon were alive today, the sight of them would make grown women leap on their chairs, clutching the skirt tails in fear. 

What of Morganucodon itself? The skeleton of M. watsoni is small (about 10 cm nose to tail base) and extremely mammal-like, with differences limited to fine anatomical details. For instance, the composition of the Moranucodon jaw-joint isn't a simple as those of true mammals, the scapula is a little bit 'reptile-like', and (according to some sources, anyway), there may have been some degree of bowing to the fore- and hindlimbs when walking or standing. This doesn't necessarily indicate sprawling limbs, but they may not have been as neatly tucked under the body as those of other mammals. Because morganucodont skeletons are so similar to those of true mammals, it seems likely that many aspects of our basic soft-tissue anatomy were established by this point of synapsid evolution - ear pinnae, fur, loose skin and so on. Direct evidence for these are currently lacking in morganucodonts, but there is indirect evidence for fur from the relatively large Morganucodon brain. Although not as large as those of later mammals, Morganucodon endocasts were expanded beyond a typical 'reptilian' condition, and much of this reflects an enlarged neocortex. This part of the brain processes sensory information, and it may be that the covering mammaliaform bodies with pressure-sensitive hairs promoted this development. I find this observation quite interesting because other lineages with fuzzy bodies - bird-like dinosaurs and pterosaurs - have also developed expanded brains and enhanced abilities to process sensory information. These enlargements are often attributed to enhanced balance and coordination, but might they also be related to the developed of sensitive fuzzy hides, as is assumed for mammals?

Because art of Mesozoic mammals scurrying about at night is so rare, I wanted to capture this in my painting. The evidence for nocturnal activity in Mesozoic mammaliaforms is not as strong as you might think - it's largely based on the (questionable?) assumption that reptiles forced early mammals into nocturnal niches and the abundance of small nocturnal mammals in the modern day - but what the heck: it makes for a fun picture. It seems nocturnal mammals are often equipped with tremendous sets of whiskers, so I put similar features on my Morganucodon to help them find their way around, They may not be 'true' whiskers, in the sense of derived, mobile whiskers of modern mammals, but it doesn't seem unreasonable to imagine long, stiffened sensory hairs of some kind developing rapidly once fur was attained.

And... blast it, I'm out of time. So many other things to say, but they'll have to wait for another time. Coming soon: various theropods, festive pterosaurs, er... and probably other things too!

Of tiny tyrants and Triassic big-heads: Tyrannosaurus rex and Garjainia madiba

This week sees two new pictures of mine being 'released' in one way or another. Much as I'd like to go into lots of detail about each, that realistically isn't going to happen anytime soon. I'm going to attempt a sort of 'picture[s] of the day'-style writing. I'm sure I can do it... right?

Chidumebi Browne's resting Tyrannosaurus teens

Two young adult old male (left) and female Tyrannosaurus on a break from pillaging and destroying the Cretaceous, distracted by a group of ruffian moths. Concept and animal colouration by Chidumebi Browne. Prints are available.

First up is one of my '£100 palaeoart offers', painted for Chidumebi Browne. Featuring Tyrannosaurus, which needs no introduction as an dinosaur most famous for antisocial tendencies, Chidumebi wanted a more relaxed approach to tyrant dinosaur art. The concept called for Tyrannosaurus at the smaller end of their size scale, settling on individuals approximating the size of the 'Jane' specimen - about half the length of a fully-grown animal. There were also requests for contrasting blue and red colours on a male and female. I was happy to oblige, seeing as some degree of dimorphism is defensible for dinosaurs even at on half their full-grown size. Like mammals and non-avian reptiles, Mesozoic dinosaurs hit sexual maturity well before attaining fully ossified, completely grown skeletons and, for Tyrannosaurus, specimens in their early teens were probably reproductively active. In that sense, some features related to sexual behaviour might be expected in 'teenage' animals. Such individuals - better considered very young adults rather than large children - look rather different to their super-size contemporaries with their longer legs and more gracile build. Some of that is obscured here by the extensive feathering covering both animals (if you look very closely, you can just make out the arms of the sitting male), but their long legs at least show through.

The concept called for a a series of moths catching the attention of the male tyrant: initially one was ordered but, even at half-size, Tyrannosaurus is pretty big, so a few more were added to make them more conspicuous. My initial thought was to use butterflies rather than moths for the role of the lepidopterans, but I was surprised to learn that butterflies don't appear in the fossil record until well after the K/Pg event. Moths have a fair, if not especially extensive Mesozoic record, so they seemed a safer bet. They certainly add an air of tranquility to the scene not featured in a lot of theropod art: well done to Chidumebi for an excellent idea.

There'll be more output from the '£100 palaeoart offers' soon, although note that the offer is now full - over-full, in fact. There's some great ideas which I'm hoping to do justice to, so thanks to all who got their orders in - the offer sold out very quickly. If you didn't manage to get something to me on time, prints are still available - wittonprints@gmail.com is the address to contact for them.

Gower et al.'s Garjainia madiba: yes, the head is that big 

Gargainia madiba sp. nov., South Africa's newest erythrosuchid. From Gower et al. 2014.

Art number 2 is a life restoration of a new species of Early Triassic stem-archosaur, the erythrosuchid Garjainia madiba, described by David Gower and colleagues in this week's PLoS ONE. Unearthed in South Africa and named for Nelson Mandela ("Mr Mandela was known affectionately as 'Madiba'" - Gower et al. 2014), G. madiba has been making surprising ripples on Twitter and Facebook because of its rather enormous head. I say surprising because, for an erythroshucid, G. madiba is fairly typically proportioned - so far as anyone can tell, anyway. We don't have anything like a complete skeleton for G. madiba, although many aspects of its anatomy are represented in fragmentary specimens. It is currently distinguished from its relatives by fine anatomical details, perhaps the most notable being its large postorbital and jugal bosses of unknown function (best seen in the reconstructed anterior aspect, above). The discovery of more substantial G. madiba fossils may reveal more obvious distinction from other erythrosuchids, but, for the time being, the best we can do reconstruction-wise is show G. prima with a madiba upgrade package. Still, given how similar the two Garjainia species seem to be, this does not seem unreasonable.

Restoring Garjainia was a lot of fun because it forced a 'back to basics' approach to the artwork where David Gower, Richard Butler and I spent a lot of time discussing proportions, muscle distribution and posture. Many fossil animals - dinosaurs, pterosaurs, etc. - have been restored so often that the basic foundations of their anatomy are very well known, but this is not so for Garjainia and other erythrosuchids. A personal revelation to come from this process was evidence for enlarged areas of axial musculature on erythrosuchid skeletons, indicated by the rather tall neural spines of their necks and backs. This might give some insight into how their large heads were supported: a particularly well-developed, strong set of axial muscles. The posterior faces of their skulls are also wide and robust, providing space sufficient to anchor powerful neck muscles. But erythrosuchid anatomy was likely not held together only by brute strength: there's also some clever biological engineering at work. Like many archosauriforms with huge-looking heads, their skulls are more gracile and lightweight than they first appear, actually being fairly narrow for much of their length and riddled with fenestrae. We tried to show the former in our anterior aspect reconstruction: note how slender the snout of the animal is compared to the cheek region. The result is a head which is undeniably large, but probably much more manageable than it first seems.

For a lot more on Garjainia and other erythrosuchids, including the life restoration in situ, full descriptions of G. madiba anatomy and revisions to the diagnosis of the group, Gower et al. (2014) can be read here (hurrah for open access!). Thanks to David and Richard for bringing me on board, and congrats to them on the paper.

Coming soon: small, brown Mesozoic mammialiaforms! Yes, they are exciting. Really.

Reference

• Gower, D.J., Hancox, P.J., Botha-Brink, J., Sennikov, A.G., & Butler, R.J. (2014) A New Species of Garjainia Ochev, 1958 (Diapsida: Archosauriformes: Erythrosuchidae) from the Early Triassic of South Africa. PLoS ONE 9(11): e111154. doi:10.1371/journal.pone.0111154

Prints, custom palaeoart, and antisocial Triceratops

Hey now - look at this: there's a way to buy high-quality prints of my artwork, and it's dead simple. Drop me an email at wittonprints@gmail.com with your requirements, pay off the invoice, and wait for your prints to arrive - hopefully within a week or so of order confirmation. Prices, sizes and all are discussed over here. Added 02/2015: visit my print store!

To celebrate this occasion, I'm also offering a limited number of über-cheap palaeoart commissions for private clients:

Yep - your own palaeoartwork, a print and delivery for just £100, which is a stupidly cheap price for original artwork. Full details (including a few important conditions) are here (edit 02/2015: not any more!). As you may expect, I can't sustain working at that price for long. For that reason, there's only five of these deals being offered, and at time of writing, three of these deals have been taken. If you want in, don't delay. Don't despair if you miss this deal but would still like your own commission: drop me a line and we might be able to work something out.

Finally, because things have been a bit quiet about here for the last month or so, here's something to fill the void: a monochrome bristly Triceratops horridus, the dromaeosaur Acheroraptor temertyorum, and an interaction inspired by the wise, yellow philosophy of The Simpsons.

"...and like people, some [animals] are just jerks."

Apologies for yet another short post. Coming soon (when I'm allowed to publish them): Mammaliaformes! Heaps of pterosaurs! Deinonychus! Diminutive tyrannosaurines! And perhaps other subjects too!