Hello there!

September 2021 update:

There’s been a major lack of activity here for a while due to a combination of computer failure and our web hosting account completely running out of disk space.


New computer has been acquired, files have been recovered, and the host has been paid for a plan upgrade, so now I can finally work on getting the backlog of the last few months’ posts uploaded.

Welcome to the long-overdue new version of Nix Illustration!

Pardon our dust – we’re working on getting everything properly set up here, and also importing in around six years’ worth of archived content from elsewhere.

(…Very, very slowly.)

Please note that unless otherwise stated, all original non-commissioned work here is published here under a Creative Commons Attribution-NonCommercial license (CC BY-NC 4.0) – you are welcome to use images for non-profit or personal purposes, provided you credit me and give proper attribution.

Please contact me via tumblr or email (mail@nixillustration.com) to enquire about custom commission work or commercial image licensing.

In the meantime, you can find more complete selections of work at any of these places:
Tumblr | Pillowfort | Twitter | Patreon

Current status:
all 2019 content uploaded
–older works in progress


Ghost pipefishes are close relatives of pipefish and seahorses, and today are represented by six different species found in shallow tropical waters of the Indo-Pacific. But while this lineage is estimated to have originated around 70 million years ago in the Late Cretaceous, their fossil record is very sparse – only three fossil representatives are currently known from the entire Cenozoic.

Calamostoma lesiniforme is one of the oldest of these, dating to the early Eocene around 50-48 million years ago. Known from the Monte Bolca fossil beds in northern Italy, it lived in a warm shallow reef environment during a time when that region of Europe was covered by the western Tethys Ocean.

Up to about 9cm long (3.5″), it was already very similar in appearance to modern ghost pipefishes, with a long tubular snout, star-shaped bony plates in its skin, two dorsal fins, and fairly large pelvic fins that formed an egg-brooding pouch in females. It probably had the same sort of lifestyle as its modern relatives, floating pointing downwards and camouflaging itself among seagrasses, algae, and corals.

One specimen preserves a small amount of color patterning, showing hints of dark banding on the pelvic and tail fins. But since modern ghost pipefish can change their coloration to better mimic their surroundings, it’s unclear whether these markings were common to all Calamostoma or were just part of this particular individual’s camouflage.


Modern birds’ upper beaks are made up mostly from skull bones called the premaxilla, but the snouts of their earlier non-avian dinosaur ancestors were instead formed by large maxilla bones.

And Falcatakely forsterae here had a very unusual combination of these features.

Living in Madagascar during the Late Cretaceous, about 70-66 million years ago, it was around 40cm long (1’4″) and was part of a diverse lineage of Mesozoic birds known as enantiornitheans. These birds had claws on their wings and usually had toothy snouts instead of beaks, and many species also had ribbon-like display feathers on their tails instead of lift-generating fans.

Falcatakely had a long tall snout very similar in shape to a modern toucan, unlike any other known Mesozoic bird, with the surface texture of the bones indicating it was also covered by a keratinous beak. But despite this very “modern” face shape the bone arrangement was still much more similar to other enantiornitheans – there was a huge toothless maxilla making up the majority of the beak, with a small tooth-bearing premaxilla at the tip.

This suggests that there was more than one potential way for early birds to evolve modern-style beaks, and there may have been much more diversity in these animals’ facial structures than previously thought.


During the Early Carboniferous, around 330 million years ago, the region that is now the East Kirkton Quarry in Scotland was located close to the equator, with a lush tropical climate and volcanic hot springs dotting the landscape. It preserves fossils of some of the earliest known fully terrestrial tetrapods, and a recent discovery shows how some of these animals were already experimenting with the shapes of their feet to better get around on land.

Termonerpeton makrydactylus is only known from a partial skeleton, and shows a mix of anatomical features that make identifying its exact evolutionary relationships rather difficult – but it was probably a very early reptilomorph, closer related to amniotes than to lissamphibians. It may also have been very closely related to the equally enigmatic Eldeceeon and Silvanerpeton from the same region, but was almsot twice their size with a estimated total length of around 70cm (2’4″).

It would have resembled a rather heavily-built lizard-like or salamander-like animal, with fairly stumpy legs and probably lacking claws on its digits. While it would have had spindle-shaped scales on its underside, and possibly small rounded scales along its sides and back, these were bony structures embedded in its skin and probably weren’t very visible externally in life.

But Termonerpeton‘s most surprising feature was its proportionally large feet with especially elongated fourth toes, which would have helped to extend its stride length for energy-efficient terrestrial locomotion and to stabilize its movement on unstable surfaces – a much more “advanced” amniote-like arrangement than expected in such an early reptilomorph, and convergently similar to to the foot shapes of some modern lizards. Its fourth toe was also unusually chunky, suggesting it may even have been bearing most of its weight on just that one digit when walking.


Euthycarcinoids were a group of arthropods that lived between the mid-Cambrian and the mid-Triassic – but despite existing for over 250 million years their fossil record is incredibly sparse, and it’s only within the last decade that they’ve been recognized as being close relatives of modern centipedes and millipedes.

The earliest members of this group were marine, living in shallow tidal waters, but they quickly specialized into brackish and freshwater habitats and were even some of the very first animals to walk on land. Fossil trackways show they were amphibious, venturing out onto mudflats to feed on microbial mats, avoid aquatic predators, and possibly lay their eggs in a similar manner to modern horseshoe crabs.

Most euthycarcinoid species are known from tropical and subtropical climates, but Antarcticarcinus pagoda here hints that these arthropods were much more widespread and diverse than previously thought. Discovered in fossil deposits in the Central Transantarctic Mountains of Antarctica, it lived in freshwater lakes during the Early Permian (~299-293 million years ago), at a time when the region was in similar polar latitudes to today with a cold icy subarctic climate.

About 8.5cm long (3.3″), it would have had a similar three-part body plan to other euthycarcinoids – with a head, a limb-bearing thorax, and a limbless abdomen ending in a tail spine – but its most distinctive feature was a pair of large wing-shaped projections on the sides of its carapace. These may have helped to stabilize its body when resting on soft muddy surfaces, spreading out its weight, or they might even have functioned as a hydrofoil generating lift while swimming.


Blesmols, or African mole-rats, are a group of rodents adapted for mole-like burrowing. Closely related to the more famous naked mole-rat, these little mammals have reduced eyes and ears along with incisors that protrude out even when their mouths are closed, allowing them to excavate tunnels primarily using their teeth.

One of the earliest known fossil blesmols is Bathyergoides neotertiarius here, from the early Miocene of Namibia about 20 million years ago. For almost a century this species was known only from teeth and partial skull remains, but recently a partial skeleton was described giving us a better idea of its overall appearance and lifestyle.

Bathyergoides was a fairly large blesmol, around 25cm long (~10″), and was already specialized for tooth-digging with a skull very similar to modern forms. It had powerful muscular forelimbs that would have been used to push back loose soil while burrowing, but unlike its living relatives it also had a long tail and relatively slender hindlimb bones – with anatomy suggesting its legs were used more for stabilizing its posture than for actively digging.

It may have had a less subterranean lifestyle than modern blesmols, digging out extensive burrows but still foraging for food above ground in a similar manner to modern semi-fossorial rodents like giant pouched rats.


Amargasaurus cazaui was a sauropod dinosaur with a very distinctive-looking skeleton, sporting a double row of long bony spines along its neck and back. It lived in what is now Argentina during the Early Cretaceous, about 129-122 million years ago, and was fairly small compared to many other sauropods, reaching about 10m in length (~33′) with a proportionally short neck compared to its body size.

And despite being known from fairly complete skeletal remains there’s still a lot we don’t know about this dinosaur – especially what was actually going on with those vertebral spines. While it’s sometimes been depicted with skin sails over the spines, for the last couple of decades the general opinion has trended towards them being more likely to have been covered by spiky keratinous horn-like sheaths.

But recently that’s been brought back into question. A detailed study of the microscopic bone structure of Amargasaurus‘ spines shows no evidence for keratin attachment and instead found textures associated with skin coverings, along with an extensive web of ligaments connecting the spines to each other along each row.

So maybe it had big flashy sails after all!

Eons Roundup 12 (& Published Art!)

It’s been a while since I last showed off some of these, but here’s some more commission work I’ve done for PBS Eons:

The metriorhynchid marine crocodilians Aggiosaurus and Cricosaurus, from “When Crocs Thrived in the Seas”

And… what’s this?

A familiar Scutellosaurus makes an appearance in a recently-published children’s dinosaur book!

April Fools 2022: The Aquatic Dinosaur That Wasn’t

So, Spinosaurus wasn’t technically the first known aquatic non-avian dinosaur.

That title instead temporarily went to Compsognathus corallestris.

While the idea that hadrosaurs and sauropods were wallowing swamp-dwellers had been completely abandoned at the start of the Dinosaur Renaissance, the new view of dinosaurs as active sophisticated animals led to a surprising aquatic hypothesis during the early days of this paleontological revolution.

A specimen of the small theropod Compsognathus discovered in southeastern France in the early 1970s was only the second skeleton ever found of this dinosaur, and came over a century after the first. It was initially thought to represent a new species since it was about 50% larger than the German specimen of Compsognathus longipes, and it seemed to have something very unusual going on with its hands – its forelimbs were somewhat poorly-preserved and distorted, and had traces of some sort of large fleshy structure around the hands that was interpreted as representing elongated three-fingered flippers used for swimming.

This wasn’t necessarily as ridiculous of an idea as it might sound. Compsognathus lived during the Late Jurassic, about 150 million years ago, at a time when Europe was a group of islands in a shallow tropical sea. A semiaquatic dinosaur specialized to swim and dive, hunting the abundant aquatic prey in its environment, and easily able to island-hop all around the European archipelago seemed at least somewhat plausible, and reconstructions of fin-handed C. corallestris even appeared in several popular dinosaur books of the time.

But it didn’t last.

Within just a few years doubt was being cast on this idea, and further studies of both known Compsognathus skeletons in the late 1970s and early 1980s concluded that C. corallestris was actually a fully-grown adult individual of the juvenile C. longipes. The French Compsognathus had normal-looking hands for its kind after all, with two large clawed fingers and a vestigial third finger, and the “flipper” impressions had just been ripples in the fossil slab.

For a long time after that the general view became that there just weren’t any aquatic non-avian dinosaurs at all – but more recent discoveries like the new Spinosaurus material and Halszkaraptor are starting to suggest that some of these animals were much more at home in the water than previously thought.

Something resembling Compsognathus corallestris might still surprise us in the future.

Retro vs Modern #23: Spinosaurus aegyptiacus

Spinosaurid teeth were first found in the 1820s in England, but were misidentified as belonging to crocodilians. It wasn’t until nearly a century later that Spinosaurus aegyptiacus was discovered and recognized as a dinosaur – and it would be another century after that before we really started to learn anything about it.


The first fossils of Spinosaurus were discovered in Egypt in the 1910s. With only a few fragments of its skeleton known it was an enigma right from the start, hinting at a large and very strange theropod dinosaur with crocodile-like teeth, an oddly-shaped lower jaw, and elongated neural spines on its vertebrae that seemed to be part of a huge sail.

A few possible extra fragments were found in the 1930s, but overall these few pieces were all that was known of Spinosaurus for a long time.

The fossils were kept in the Paleontological Museum in Munich, Germany,a building that was severely damaged during a bombing raid in World War II. Many important specimens were destroyed, including Spinosaurus, and only the published drawings and descriptions of the bones remained.

So for the next several decades Spinosaurus remained a very poorly-understood mystery. During this period it was generally depicted as a generic “carnosaur“, often modeled on something like Megalosaurus, in the standard-for-the-time tripod pose and with a Dimetrodon-like sail on its back.

Interestingly a 1930s skeletal reconstruction shows Spinosaurus with an unusually long torso and fairly short legs, details that are surprisingly modern despite the retro posture.


In the 1980s some partial snout bones from Niger were recognized as having similarities with the jaw of Spinosaurus. Around the same time the fairly complete skeleton of Baryonyx was discovered, and along with further spinosaurid discoveries in the mid-to-late 1990s a decent idea of what Spinosaurus might have looked like began to emerge.

It was reconstructed with a long kinked crocodilian-like snout, a ridged bony crest in front of its eyes, an S-curved neck, and large thumb claws on its hands – an interpretation that was heavily popularized by Jurassic Park III in the early 2000s, bringing this enigmatic dinosaur to public attention and portraying it as a fearsome super-predator bigger than Tyrannosaurus.


Despite attempts to locate more complete Spinosaurus remains, only fragments continued to be found, and it remained a frustratingly poorly-known species even into the early 2010s.

Finally, in 2014, almost a full century after it was first described and named, Spinosaurus started to reveal its secrets with the announcement of the discovery of the most complete skeleton so far, discovered in the Kem Kem fossil beds in Morocco. Its body was still only partially represented, but it included skull fragments, part of a hand, a complete leg and pelvis, some sail spines, and several vertebrae from the neck, back, and tail.

And nobody was expecting what these pieces revealed.

It had a very long torso and proportionally short stumpy legs, and was reconstructed with a huge distinctive “M-shaped” sail on its back. Its feet had flat-bottomed claws and its “dewclaw” toe was enlarged into an extra weight-bearing digit – adaptations for spreading its weight over soft muddy ground, and suggesting its feet may also have been webbed. Initially it was also presented as possibly being quadrupedal, due to how far forward its center of mass seemed to be, reviving an odd idea from the late 20th century.

Along with its long crocodile-like head and conical teeth, this was interpreted as evidence it was a semiaquatic fish-eating swimming animal – potentially making it the first known semiaquatic non-avian dinosaur. Spinosaurids had been suggested to be specialized piscivores before, especially since Baryonyx had been found with fish scales in its stomach, but they were generally assumed to be more like modern grizzly bears, wading into water to hunt but not being habitual swimmers. Spinosaurus’ weird croco-duck proportions, however, seemed like they might be much more suited to watery habitats than to the land.

Since Spinosaurus had become a popular dinosaur with the general public by that point, the discovery was big news – and a big controversy for a while. It was so bizarre that some paleontologists were skeptical of the radical new interpretation, wondering if the measurements of the skeleton were correct or if the short legs were even from the same individual or the same species as the rest of the bones.

After a while the new proportions were accepted as fairly accurate, and over the next few years attention turned to instead figuring out just how this animal worked and how aquatic it actually was. An earlier isotope analysis of its teeth supported a semiaquatic lifestyle similar to crocodiles and turtles, but a buoyancy study argued that it might not have been able to dive below the water suface and its sail made floating unstable – but also found that its center of mass was closer to its hips than previously calculated, suggesting it could walk bipedally after all.

Then in 2020 came another surprise: more of the tail of the new specimen had been found, and it was just as weird as the rest of Spinosaurus. Its tail was a huge vertically flattened paddle-like fin supported by long thin neural spines and chevrons, resembling a giant eel or newt more than a dinosaur and also giving some more weight to the idea that it was a swimmer.

Our modern view of Spinosaurus is still evolving, and likely to be full of even more surprises in the future as we discover more about this unique dinosaur. But we at least know it lived in what is now North Africa during the Late Cretaceous, about 99-93 million years ago, and whether it was a swimmer or wading generalist predator it was one of the largest known theropods to ever live, estimated to have reached around 16m long (~52ft).

While the “M-shaped” sail reconstruction has been popularized by the recent discoveries, the exact shape of this structure is still unknown. Like with other sailbacked animals it’s also not clear what it was for, with ideas including temperature regulation, visual display, supporting a fatty hump, and a potential hydrodynamic adaptation.

EDIT: And while I was working on this entry (late March 2022) I missed that another study had just come out with more anatomical support for swimming Spinosaurus!

Retro vs Modern #22: Tyrannosaurus rex

Probably the most famous and popular dinosaur of all time, Tyrannosaurus rex is also the only species commonly known by both its full scientific name and its abbreviation T. rex.


Fragments of what we now know are Tyrannosaurus fossils were first found in the Western United States in the 1870s, but it wasn’t until the early 1900s that a couple of partial skeletons were discovered and recognized as belonging to a new species of huge carnivorous theropod.

With its charismatic and memorable name meaning “tyrant lizard king” it was an immediate hit with the general public, portrayed as the “king of the dinosaurs” in pop culture and as the dramatic nemesis of Triceratops.

Like other bipedal dinosaurs of the time it was depicted in an upright kangaroo-like tripod pose, cold-blooded and lizard-like. Sometimes it was also shown with three-fingered hands, since its arms were poorly known for a long time – and while the closely-related tyrannosaurid Gorgosaurus was known to have had just two fingers, this wasn’t confirmed for Tyrannosaurus until the late 1980s.


During the 1970s and the early Dinosaur Renaissance it became obvious that bipedal dinosaurs like Tyrannosaurus couldn’t have stood so sharply upright without dislocating their hips and tail vertebrae.

Jurassic Park was influential in introducing this new corrected posture to the general public in the early 1990s, presenting a powerful and active predator with a more bird-like horizontal stance and its tail acting as a counterbalance. Reconstructions inspired by this portrayal became standard for Tyrannosaurus during the 1990s (although the old-style tripod remains in public consciousness even decades later), and while it didn’t tend to get as heavily shrinkwrapped as some other species it was still common for a while to push its belly ribs in as much as possible to make its bulky body look skinnier and more “athletic”.

The 1980s and 1990s were also a time when discoveries of new Tyrannosaurus specimens began to become much more frequent, improving knowledge of the species’ anatomy, biology, and life history. Some, like Sue, Stan, and the Dueling Dinosaurs, would also unfortunately end up becoming highly controversial, tied up in legal disputes for years and sold for multi-million-dollar prices.


After the explosion of feathered non-avian dinosaur specimens from China in the mid-1990s, eventually the small feathered tyrannosauroid Dilong was discovered in the early 2000s, followed by the much larger-bodied Yutyrannus in the early 2010s.

While these tyrannosauroids weren’t particularly closely related to Tyrannosaurus itself, the question of potential tyrant fuzz still began to loom, and for a while in the 2010s highly fluffy T. rex interpretations were popular in paleoart. But in the late 2010s a study of known skin impressions from Tyrannosaurus and several of its closer relatives showed that small pebbly scales were known from various parts of the body, and suggested that these particular dinosaurs were most likely primarily scaly. Sparser fluff was still possible on parts of the body, however, similar to the hair on modern elephants, and it’s also possible that juveniles were much fuzzier.

(While this is disappointing for fans of huge fatbird T. rex, it’s also a great example of the scientific process. The skin impressions hadn’t ever been properly described before this point, and the scaly interpretation had mostly been an assumption. Speculative fluffiness prompted all the skin evidence to actually be consolidated, and now we know a lot more about Tyrannosaurus’ potential outward appearance than we did before.)

Arguments about lips in theropod dinosaurs also went back and forth during the 2010s, with interpretations ranging from tight-skinned crocodilian-like snouts with exposed teeth to fleshy lizard-like lips similar to modern Komodo dragons. There’s not really a consensus yet, but since most non-beaked tetrapods do have lips the safe bet is still that dinosaurs like Tyrannosaurus would have had them too. 

Our modern view of Tyrannosaurus is a much chunkier animal than older interpretations, with  more extensive soft tissue, properly-positioned belly ribs showing that it had a barrel-shaped pot-bellied body, and its tail being thicker-muscled than previously thought.

Living across western North America – then the island continent of Laramidia – during the very end of the Cretaceous, about 68-66 million years ago, it was one of the largest terrestrial carnivores to ever live. The very biggest known specimens are estimated to have been as much as 13m long (~43′), with the proportionally large head making up around 1.5m of that (~5′).

Its skull was boxy at the back but narrow along the snout, allowing its forward-facing eyes to have hawk-like stereoscopic vision. Large fenestrae and a “honeycomb” of air spaces reduced the weight of the skull, while reinforced fused bones strengthened it, and Tyrannosaurus is estimated to have had an incredibly powerful bone-crushing bite force.

It had a highly developed sense of smell, and its hearing was geared towards low-frequency sounds. The texture of its skull bones suggests it may also have had thick toughened keratinous skin and bumps over its face, which might have been involved in head-shoving and headbutting behaviors.

Although proportionally tiny for its overall size, its arms were still rather beefy, with large areas for muscle attachment with “meathook” claws that may have been used to hold onto struggling prey.

As a heavily-built bulky carnivore it probably wasn’t especially fast, and its legs were adapted for energy-efficient walking rather than running. It may have been a long-distance stalker, only using short bursts of speed in a final ambush – and like most large carnivores it would have also opportunistically scavenged on carcasses, too.

Specimens once argued to belong to a separate smaller species of tyrannosaur, Nanotyrannus, are now generally accepted as actually being juvenile Tyrannosaurus. They show a surprising amount of physical change as these animals aged, starting out leaner-built with longer legs more suited for speed, slender more delicate snouts, and only developing the characteristic chunky adult proportions during a huge growth spurt in their mid-to-late teens.

Meanwhile, the latest big controversy over this dinosaur as of March 2022 (because there’s always something) is a study proposing splitting Tyrannosaurus into three separate species: T. rex, T. imperator, and T. regina. It doesn’t seem to be going down well, but much like the feather situation it probably at least means we’re going to see a lot of further investigation over the next few years.