Attempted adventures in dinosaur facial restoration, starring Giganotosaurus

Giganotosaurus carolinii carries the remains of a juvenile rebacchisaurid while sporting a bunch of crazy tissues on its face. Big scales, horns, and is that some kind of thick pad over its snout? Is all this artistic speculation or something inferred from fossils? Read on...

Up until now, my palaeoart career has not crossed paths with carcharodontosaurids, the gigantic, charismatic and famous allosauroids best known for Acrocanthosaurus, Giganotosaurus, Carcharodontosaurus and, most recently, Meraxes. This month, however, I finally had cause to restore Giganotosaurus carolinii, the largest of the group and, potentially, the largest of all theropods.

Carcharodontosaurids are, at first glance, not too challenging to restore: take an Allosaurus, turn everything up to 11 and job done, right? Well, maybe not. Not only are the proportions of carcharodontosaurids (and, to be fair, carcharodontosaurians in general) subtly different from their allosauroid ancestors, but their jaws and eye regions are characterised by a suite of complex sculpting and rugosities. It’s thought that these are epidermal correlates (Sereno and Brusatte 2008): distinctive bone surfaces and histological patterns that record different skin types interacting with the underlying bone (Hieronymus et al. 2009). I’ve written quite a lot about epidermal correlates at this blog because they provide heaps of important external soft-tissue information without direct soft-tissue fossilisation and learning to spot them, in my view, is an essential skill for any budding palaeoartist.

The presence of epidermal correlates on carcharodontosaurid skulls means that we can’t take an “anything goes” approach to restoring Giganotosaurus or its close relatives; instead, there probably is a “right”, or at least "more defensible", way to approach depictions of their faces. Alas, to my knowledge, no specific investigation has been conducted into what carcharodontosaurid skull textures represent despite our interest in other dinosaur epidermal correlates (e.g. Hieronymus et al. 2009; Carr et al. 2017; Delcourt 2018). This means there’s not yet a “go-to” study to provide artists with answers for restoring these animals and anyone wanting to illustrate Giganotosaurus credibly has to make their own interpretations from descriptions and illustrations in scientific literature. Having just been through this process myself, and realising that Giganotosaurus is a fan-favourite, I thought it might be of interest to share my thoughts here. I want to be upfront by declaring that the following deductions are little more than best guesses; without having direct experience of Giganotosaurus fossils I can’t write anything definitive about what Giganotosaurus looked like. Think of the following more as a discussion piece than a rigorous guide, and I welcome input and insight from others if I’ve made errors.

As I understand it, most of what's been illustrated of Giganotosaurus is shown above in this compilation of figures from Coria and Salgado (1995; greyscale graphics) and Novas et al. (2013; colour). Not as much as you might expect, right? There's a lot more known of this species that hasn't been published yet, some of which contain crucial information for palaeoartists (and I guess for scientists too).

And it’s in this spirit that, right off the bat, we need to mention that researching Giganotosaurus is pretty challenging. Its fossils are thinly documented despite Giganotosaurus being one of the more completely known carcharodontosaurids and, even today, almost 30 years since it was announced to the world, we only have a fraction more information available to us than when it was first named in 1995. Just a handful of its bones have been figured so good photos or illustrations of several fossils relevant to this conversation have not been published (Coria and Salgado 1995; Coria & Currie 2002; Novas et al. 2013). Thus, anyone trying to restore this animal from scientific papers alone will struggle for information and a lot of secondary sources — online photographs of fossils and casts, skeletal reconstructions and museum mounts etc. — are essential to obtaining basic information about its proportions and size, even if they risk introducing reconstruction errors. Closely related taxa and comparative descriptions (i.e. “Mapusaurus has a more rugose snout than Giganotosaurus”) are critical too, providing crucial details not mentioned in dedicated Giganotosaurus papers. I mention this because it means that, from the get-go, we’re not in an ideal research scenario for a palaeoartwork, and this makes the possibility of errors in interpretation all the greater.

With appropriate caveats established, let’s dive into this discussion. As with most theropods that have rugose, textured faces, our attention here is going to be on the bones of the snout and orbital region, as these are the principal areas to bear features that might signify epidermal tissues. The carcharodontosaurid fossil record contains a large number of maxilla bones (the main tooth-bearing bone of the upper jaw) and this is good news for artists, as the lateral surfaces of these potentially tell us a lot about the skin on the side of the upper jaw. The typical maxillary rugosity for carcharodontosaurids is well documented across several species, especially Carcharodontosaurus saharicus, Eocarcharia, Mapusaurus and Meraxes. It comprises a series of sub-vertical grooves and pits (Stromer 1936; Sereno et al. 1996; Coria and Currie 2006; Brusatte and Sereno 2007; Sereno and Brusatte 2008; Canale et al. 2022) and some species (e.g. C. saharicus, Eocarcharia, Mapusaurus) supplement these with prominent ridges extending along the base of the antorbital region. These bars separate the rugose maxillary body from the smoother bone of the antorbital fossa: that slightly impressed region of bone surrounding the antorbital fenestra.

The maxilla of Eocarcharia dinops, as illustrated by Brusatte and Sereno (2008), shows the texturing typical of carcharodontosaurid snouts. Note the ridge dividing the textured region from the smoother antorbital fossa: this feature isn't seen in all carcharodontosaurids but might tell us something about skin types all the same. The fossa region is particularly big in this species.

The texturing characterising carcharodontosaurid maxillae may be somewhat less pronounced in Giganotosaurus and thus, perhaps like Acrocanthosaurus, its maxillae may have been on the smoother end of the rugosity scale (Coria and Currie 2006; Eddy and Clarke 2011; Novas et al. 2013). All else being equal, this might imply differences in facial anatomy within Carcharodontosauridae: whatever those grooves and pits signify may not have been as exaggerated in some species as others. A caveat here is that, as is often the case with skin-altered bones, larger carcharodontosaurid individuals tend to have more exaggerated rugosity profiles than smaller ones (Coria and Currie 2006; Canale et al. 2014), suggesting a link with body size or age as well as differences between species. We probably want a number of maxillae from a range of differently aged individuals to establish whether a species has consistently smoother jawbones than its relatives.

Comparing these maxillary features with existing interpretations of dinosaur epidermal correlates provides potential insights into their significance. The textures in question are often likened to those adorning the lateral surfaces of abelisaur skulls and, if so, we might follow Delcourt (2018) in inferring that they represent scale correlates. This seems sensible to me, certainly more than other possible jaw coverings. Carcharodontosaurid maxillae lack the projecting rugosities consistent with armoured dermis or the branching neurovascular channels and oblique foramina found under beak or horn tissues (Hieronymus et al. 2009). Furthermore, the ridges bordering the antorbital regions in some carcharodontosaurids are inconsistent with beaks: cornified sheath tissues tend to terminate with obvious steps downwards into smoother neighbouring bone, not upwards to ridges of rugose bone (Hieronymus et al. 2009).

The skull of a common snapping turtle Chelydra serpentina. The more rugose parts of this skull correspond to regions covered in large scales, while the slightly finer rugosity around the jaws demark the distribution of the beak (also note the stepped topography at the beak/scale transition). Prominent ridges occur around the eyes and nose where large scales meet softer tissues: perhaps this is analogous to what we're seeing in those ridged carcharodontosaurid maxillae?

Raised bony ridges are seen, however, around the skull openings of reptiles with tough, tightly-adhering facial skin like crocodylians and certain turtles, marking some boundaries between thick, relatively immobile skin and softer, more flexible regions. We might expect the antorbital skin of theropods to flex slightly during breathing, as it does in birds, and I wonder if we're picking up some evidence of that in carcharodontosaurids? The notion that carcharodontosaurid maxillary skin might be tough and immobile is not without precedent, as early members of the broader Carcharodontosauria clade are thought to have had maxillary skin of this nature (Barker et al. 2019). If there really was a distinction in skin flexibility in the snouts of these animals it may have been obvious in life, as it is in crocodylians and turtles (I realise this sounds like advocating some form of shrinkwrapping — lightning flashes in the distance, thunder rumbles — but we can't overlook the fact that osteological features do, sometimes, correlate with skin types in living animals). I took these reptiles as inspiration in my reconstruction, giving Giganotosaurus a series of large, thick scales over the side of its upper jaw that terminate sharply around the antorbital region. I retained a full set of lips for reasons that have been thrashed out too many times to bear repeating here, except to mention that — like those of tyrannosaurs — carcharodontosaurid maxillae seem to constrain their rugosity to regions above the toothrow, suggesting whatever skin anchored above the labial foramina (the row of perforations along the jaw) was not so tightly anchored next to the teeth.

Immediately above the maxillae are another set of sculpted bones: the nasals and lacrimals. Collectively, these bones form the various fins and crests that line the top of the snouts in many allosauroids (see Chure and Loewen 2020 for a great visual of these), but the carcharodontosaurid condition is not typical of this wider clade. Nasal material is known for Giganotosaurus but it was not featured in its original description, nor (to my knowledge) has it been illustrated elsewhere. What’s hinted at in various reconstructions and papers is that Giganotosaurus joins Mapusaurus, Meraxes and Carcharodontosaurus in having especially sculpted nasal bones over the maxillary region, specifically bearing deep, generally parallel-sided grooves crossing transversely over the dorsal surface and vertically on the lateral face (Coria and Currie 2006).

The right nasal of Mapusaurus rosae, one of the better-illustrated examples of the crazy rugosities developed on these bones by some carcharodontosaurids. A shows the lateral view, B is dorsal, from Coria and Currie (2006).

These bones cannot be described as forming narrow crests as they can for Allosaurus and kin because their texturing meets in the middle of the skull and they are not pinched into long, narrow fins (Sereno et al. 1996; Coria and Currie 2006). Accordingly, some common artistic interpretations of these structures as supporting crests or a series of hornets over carcharodontosaurid faces (which I first assumed when embarking on this painting project, I think incorrectly: see below) may be erroneous: whatever skin made these features extended over the entire dorsal surface of the nasals as well as across the upper lateral region of the snout. Exactly what’s happening here is unusual among theropods, but the rugosity depth almost certainly implies some extensive cornificiation. I'll go further to say that, to the best of my knowledge, deep, subparallel grooves are uniquely associated with cornified pads growing at shallow angles to the underlying bone (Hieronymus et al. 2009). If correct, might we infer that heavy, thick bars of densely keratinised tissue adorned the top of carcharodontosaurid skulls? Cornified pads are predicted in this region elsewhere within Theropoda (e.g. within abelisaurids: Delcourt 2018) so such a suggestion isn’t entirely without precedent, but I'm not sure we've viewed carcharodontosaurids with such heavy ornament before. It would be great to see some actual research on this to investigate what’s really going on with these bones. Giganotosaurus striding around with fat cornified pads atop its face would be all sorts of awesome, especially given that we already think other regions of carcharodontosaurid faces might be adapted for headbutting (e.g. Sereno and Brusatte 2008; Cau et al. 2013).

An earlier version of my Giganotosaurus reconstruction with individual hornlets above the snout: thinking again on this topic, I probably got this wrong as the nasal texturing isn't consistent with the bones that underly hornlets in living species. I've already sent myself to bed without dinner as punishment.

The corrugated nasals are bordered posteriorly by further rugosities around the orbit. This is actually one of the better-known parts of the Giganotosaurus skull and it has been illustrated (Coria and Salgado 1995) so we can be pretty confident about what this region generally looked like, even if a lack of a comprehensive description means it’s difficult to know exactly what sort of rugosities it bears. In terms of basic structure, a rounded, horn-shaped process sits atop the lacrimal (the bone in front of the orbit) and a prominent boss projects above and somewhat laterally from the postorbital (the bone behind the eye). As seems typical for carcharodontosaurids and, indeed, for carcharodontosaurians in general, the latter slopes back and downward somewhat such that Giganotosaurus and kin probably looked perpetually worried, their postorbital bosses creating the appearance of a furrowed brow. 

From what I can gather, the ultra-rugosity of the nasal bones doesn’t extend fully over the eyes in Giganotosaurus or its relatives. I suspect, based on what we see in better-illustrated carcharodontosaurids, that this reflects adornment of the lacrimal process with a cornified sheath rather than a pad. This creates the potential for a sharper horn than implied by the underlying bone shape, although it just as easily could be an exaggeration of the relatively blunt underlying bone structure. As is widely known in palaeoart circles, it can be difficult to predict the exact shapes cornified sheaths will take, even in modern species (Angst et al. 2020).

The postorbital boss variation of carcharodontosaurids, as illustrated by Sereno and Brusatte (2008): A shows the simpler morphology of Eocarcharia dinops; B shows Carcharodontosaurus saharicus.

Similar textures seem to have extended continuously onto the postorbital boss in derived carcharodontosaurids, such that we might imagine a continuation of the sheathed skin of the lacrimal onto this region (Coria and Currie 2006; Sereno and Brusatte 2008; Canale et al. 2022). There is, however, some variation of boss morphology within the clade in that some species have relatively smooth, rounded bosses (e.g. Sereno and Brusatte 2008; Cau et al. 2012): this is another area where more information specifically on Giganotosaurus would be welcome. For those species lacking pronounced texturing, I wonder if we’re dealing with big scale correlates rather than a surface covered with thick, densely keratinised tissue? These skin types may not be mutually exclusive however, as there is precedent for scale correlates showing signs of cornification in some dinosaurs (Hieronymus et al. 2009). A scaly postorbital boss in a young animal could well develop into a more cornified, horny structure in an adult. Again, more specimens of different growth stages might be needed here to be certain of true differences between species.

The result of all this noodling: Giganotosaurus looking a little more knobbly than usual, and also a bit world-weary thanks to that postorbital boss. Maybe the pressure of the "who's the biggest theropod" competition is pretty intense for these guys.

Putting all this together resulted in the image of Giganotosaurus that accompanies this post. Thanks especially to the big cornified pad bridging the middle of the skull, this is a face that looks more heavy-duty than we’re used to and maybe less generically “allosaurian”. But unfamiliar as it is, I’m happy with this outcome because following evidence to unexpected results is one of the great joys of palaeoart, and I always enjoy rationalising an unusual reconstruction from a foundation in science rather than mere speculation. But, again, I want to stress that this is just my interpretation of information gleaned from a less-than-ideal representation of Giganotosaurus in technical literature. This means I may have made errors obvious to those more experienced with these fossils and, moreover, when the structures discussed here are finally studied for their soft-tissue significance, the outcomes may be very different.

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References

• Barker, C. T., Naish, D., Newham, E., Katsamenis, O. L., & Dyke, G. (2017). Complex neuroanatomy in the rostrum of the Isle of Wight theropod Neovenator salerii. Scientific Reports, 7(1), 1-8.
• Brusatte, S. L., & Sereno, P. C. (2007). A new species of Carcharodontosaurus (Dinosauria: Theropoda) from the Cenomanian of Niger and a revision of the genus. Journal of Vertebrate Paleontology, 27(4), 902-916.
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• Canale, J. I., Apesteguía, S., Gallina, P. A., Mitchell, J., Smith, N. D., Cullen, T. M., ... & Makovicky, P. J. (2022). New giant carnivorous dinosaur reveals convergent evolutionary trends in theropod arm reduction. Current Biology.
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• Coria, R. A., & Currie, P. J. (2003). The braincase of Giganotosaurus carolinii (Dinosauria: Theropoda) from the upper cretaceous of Argentina. Journal of Vertebrate Paleontology, 22(4), 802-811.
• Chure, D. J., & Loewen, M. A. (2020). Cranial anatomy of Allosaurus jimmadseni, a new species from the lower part of the Morrison Formation (Upper Jurassic) of Western North America. PeerJ, 8, e7803.
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• Coria, R. A., & Salgado, L. (1995). A new giant carnivorous dinosaur from the Cretaceous of Patagonia. Nature, 377(6546), 224-226.
• Delcourt, R. (2018). Ceratosaur palaeobiology: new insights on evolution and ecology of the southern rulers. Scientific reports, 8(1), 1-12.
• Eddy, D. R., & Clarke, J. A. (2011). New information on the cranial anatomy of Acrocanthosaurus atokensis and its implications for the phylogeny of Allosauroidea (Dinosauria: Theropoda). PloS one, 6(3), e17932.
• Hieronymus, T. L., Witmer, L. M., Tanke, D. H., & Currie, P. J. (2009). The facial integument of centrosaurine ceratopsids: morphological and histological correlates of novel skin structures. The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology: Advances in Integrative Anatomy and Evolutionary Biology, 292(9), 1370-1396.
• Novas, F. E., Agnolín, F. L., Ezcurra, M. D., Porfiri, J., & Canale, J. I. (2013). Evolution of the carnivorous dinosaurs during the Cretaceous: the evidence from Patagonia. Cretaceous Research, 45, 174-215.
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Can studies of living animal colour constrain the colours of dinosaurs? A case study with big theropods

Mighty Tarbosaurus bataar carries its Therizinosaurus cheloniformis prey through a Masstrichtian forest in the rain. Colour-wise, I've decided that a ruddy-orange dorsum, light venter, disruptive black stripes, rings and large eye masks are meant to hide this 10 m long animal during predatory acts. But without direct palaeocolour data for Tarbosaurus, could we ever evaluate how sensible this colour scheme is?

Today we’re looking at one of the most commonly asked questions about restoring extinct dinosaur appearance: colour. For centuries, queries about the colours and patterns of dinosaurs, and, indeed, most extinct vertebrates, have been effectively non-answerable, save for some arm waving about the merits of camouflage for predation and display patterns for social signalling. Nowadays, advances in analyses and understanding of fossil pigments have allowed us to reconstruct the foundation colours of several dinosaurs in detail, along with those of other popular taxa like pterosaurs and marine reptiles (see Vinther 2015 and Smithwick and Vinther 2020 for overviews). This new frontier in dinosaur science has helped to flesh out not only the life appearance of dinosaurs, but also their ecology: their habitat preferences, their daily activity patterns, their predation concern and so on (e.g. Vinther et al. 2016).

Deducing dinosaur colour to this level of precision requires exceptionally high-quality preservation of their skin, down to the microscopic level, so that their pigment cells (melanosomes) can be identified. Unfortunately, this excludes the vast majority of dinosaur specimens from such analyses. Dinosaur skin is not only rare, but often occurs as mere sediment impressions rather than films of geochemically-preserved organic matter. This preservation style applies to a great number of the most famous dinosaurs so, unless some radical new science finds a way to assess colour from skin texture alone, the colours of our favourite extinct saurian taxa will probably be lost to time forever.

But can we tackle this problem from another angle? In recent decades, biologists have made enormous strides in understanding living animal colouration, looking at how it relates to habitat preferences, camouflage, signalling behaviour, body size, posture, visuality acuity and so on. Has the science around modern animal colouration advanced to the point where we can start to make tighter predictions about the colours of extinct animals? I regard this as an important question because, as much as our depictions of dinosaur anatomy have tightened since the late 20th century, our application of colour is still pretty lawless, even among professional palaeoartists. We present the same animals with colour schemes that are totally adaptively opposed to one another — one artist’s vivid blue hadrosaur is met with another’s muted browns and reds — and yet they’re both meant to be of equal scientific credibility. But how can that be so? Colours and patterns are generally thought to be under the same adaptive pressures as other parts of animal anatomy and thus should correlate, to a greater or less extent, with aspects of behaviour and ecology. There probably is, at some level, a "right" and "wrong", or at least a "likely" and "less likely" aspect to colour restoration, just as there is with all other aspects of palaeoartistry. But how can we evaluate this without palaeocolour data? Enter, stage left, the last two decades of studies of living animal colour. Can they help constrain, even in a general way, our efforts at colouring animals from Deep Time?

What flavour Australovenator wintonensis is your favourite — red, green, blue or orange? It's strange that, as consistent as we're getting with depicting dinosaur anatomy, you could present any one of these contrasting colour variants with equal scientific validity, even though they each imply very different interpretations of Australovenator biology.

Time for a case study

To investigate this, I thought we could look at a well-known group of dinosaurs to see what, if anything, living animals might suggest about their colouration. As you’ve guessed from the article title, we're using big predatory theropods for this exercise, for several reasons: 1) they’re a popular art subject, so this article should be of wide interest; 2) as regular readers will have worked out, I’m currently involved in a few big theropod projects so have been drawing them fairly continuously for a while now; and 3), the biology and ecology of big theropods are comparatively well-researched, and that helps when plugging fossil data into models of extant animal colour. And yes, we could restrict this to a more specific theropod clade but, from what I know about giant predatory dinosaurs, I’m not sure the conclusions we’d draw for big allosauroids would be much different to those of tyrannosaurines or large megalosauroids. If we’re sticking to what we know about these animals, not what we speculate and imagine about them, they only offer so much data to compare against living species.

There are plenty of caveats with this comparison, of course. No living creature is ecologically or phylogenetically close to the largest Mesozoic theropods, and our modern environments are different to those inhabited by our case study subjects. But we might also consider the importance of uniformitarianism, the adage that “the present is the key to the past”. We can’t say whether modern animals are perfect models for the colour of Mesozoic species, but they offer the only large, statistically-viable sample size of biological colour for us to work with. We are surely better off making informed guesses about extinct animal appearance using modern species as a guide, dodging known pitfalls where we can, than simply speculating wildly.

More worrying than concerns about comparing the past with today is that the controlling factors of animal colouration are extremely complicated, and it’s not clear how we can account for this. Indeed, for all of our science and ideas around animal colour, we still have lots to learn about it. Many popular, widely communicated interpretations of animal colours and patterns are only now being experimentally evaluated (Caro 2005), which means we are still struggling to understand some foundational aspects of certain colour schemes (Caro 2013). This is especially the case for predatory species, the colours of which have been relatively unexplored compared to those of prey animals (Pembury Smith and Ruxton 2020). To that end, we must temper our expectations. As neat as it would be to pour details like extinct animal size, habitat preference and trophic level into an algorithm to receive — *ping!* — a series of likely colours and patterns, our conclusions here, if any, are going to be of a more generalistic, broader nature.

Camouflage and detectability in large living predators: what does it mean for big theropods?

For all the new work that’s been done on animal colour, we still recognise that the principal pressures on animal colouration are essentially what Darwin observed in his 1871 book The Descent of Man. This is a conflict between natural selection, which promotes colour configurations that help animals remain undetected by predators, avoid temperature stress and generally survive from day to day, and sexual selection, which promotes the adoption of bold, broadcasting colours and patterns that attract mates and deter social rivals. So the first thing we might explore for big Mesozoic dinosaur predators is how our largest living terrestrial carnivores express this conflict: are they more concerned with basic natural functionality or sexual signalling? We're specifically interested in our giant theropod ecological analogues here: big animals that hunt and kill relatively large prey items. Predators that subsist on smaller, bite-sized animals don't qualify, because their ecology isn't sufficiently similar.

A selection of the largest predatory animals of modern times and their camouflage schemes, universally showing a strong adaptive emphasis on concealment regardless of habitat type, phylogeny or locomotor method. A, Ora, or Komodo dragon Varanus komodoensis (background matching); B, lion, Panthera leo (background matching); C, tiger Panthera tigris (disruptive colouration); D, polar bear Ursus maritimus (background matching); E, saltwater crocodile Crocodylus porosus (background matching); F, golden eagle Aquila chrysaetos (background matching); G, great white shark Carcharodon carcharias (countershading). All images from Wikimedia: A, Yuliseperi2020, CC BY-SA 4.0; B, Bernard DUPONT from FRANCE, CC BY-SA 2.0; C, Charles J. Sharp, CC BY-SA 4.0; D, Andreas Weith, CC BY-SA 4.0; E, fvanrenterghem, CC BY-SA 2.0; G, Juan Lacruz, CC BY-SA 3.0.

Across vertebrate groups, and across habitat types, our biggest modern predators are pretty consistently (maybe entirely consistently) primarily coloured for concealment: that is, they have camouflaging colours and patterns which hide their presence from their prey. This applies as much to mammals, which are a relatively drab group overall on account of several ecological and physiological factors (Caro 2013), as it does to clades that have the adaptive capacity to produce the most brilliant and striking colour schemes in nature, such as lizards, snakes, birds and fish. So maybe that’s our first note: big predators in the modern day are all about cryptic colouration, with little in the way of conspicuous display patterning.

Research on the impact of body size on predatory ecologies sheds light on why big predators seem to be consistently camouflage-coloured, and it’s a simple explanation: bigger animals are generally more conspicuous than smaller ones, even when they're trying their best not to be seen. The relationship between predator size and concealment capacity is still being investigated but a trend between size and conspicuousness seems to apply widely across Animalia, even in species with famously adept camouflage adaptations, like chameleons (Cuadrado et al. 2001; Pembury Smith and Ruxton 2020). Size doesn’t just affect detectability, either: it also correlates with prey response. Bigger predators instigate more vigorous reactions than smaller ones, such that prey species react sooner, flee further, or initiate more aggressive counter-responses (Stankowich and Blumstein 2005). There are strong pressures, therefore, on big predators to do what they can to remain hidden. Their size already puts them at a disadvantage for stealthily approaching prey, and they are going to have to run further or fight harder once they give up their hiding spot. Given that the largest theropods are the biggest terrestrial predators that have ever lived, we have to wonder what this link between body size and cryptic capacity implies for their colouration. Is one obvious inference that big theropods needed all the help they could get to remain inconspicuous? Would predators already handicapped by their greater detectability and exaggerated prey responses really have some of the signalling-dominant, hyper-obvious colour schemes we've given them from time to time?

Giganotosaurus adapted for the open county with high-bodied, sharply marked countershading, from my recent post about the possible facial anatomy of this animal. But note the ornament on this animal's head: I feel I gave it a pretty meaty set of soft-tissues around its snout and eye, but Giganotosaurus is still pretty undecorated compared to some theropods. Is this something we can read into — does the extent of cranial ornament tell us something facial colouration?

While fossils do not tell us anything about this correlation directly, I wonder if some anatomical evidence points to larger predatory dinosaurs aiming to be less conspicuous. Mid-and large-sized theropod fossils tend to have bony cranial ornaments more often than smaller ones (Gates et al. 2016), but in my estimation (by which I mean, this hasn't been verified by any study), the ornamentation in very large species is generally reduced and less spectacular than that of their smaller cousins. In tyrannosauroids, for instance, we see a general shift away from tall midline cranial crests in smaller, earlier species towards low-relief rugose surfaces, small horns or blunt bosses in larger taxa (Gates et al. 2016). Indeed, the very largest theropods are some of the dullest-looking, at least in terms of cranial ornament. Consider the flattened orbital bosses and rostral rugosities of Tyrannosaurus and Tarbosaurus, or the low, corrugated textures over the snouts of giant carcharodontosaurids. We can only speculate on what impact these ornaments might have had on theropod camouflaging efforts, but it’s well-established that distinctive body outlines can increase detectability, to the extent that modern predators attempt to hide them from their prey where possible (see below).

Whatever their adaptive significance, these reduced facial ornaments give us grounds to think about cranial colouring. Faces are often sites for signalling patterns and colours in modern species (e.g. Caro et al. 2017) and a reduction in bony facial ornament could indicate a lessened emphasis on this behaviour, possibly including muted facial colouration. A caveat here is that elaborate osteological features are only ever suggestive of striking colours and patterns, not directly correlated. But part of the palaeoart game is looking for clues about the nature of these animals wherever we can, and an absence or reduction of showy features is something we can factor into the reasoned speculation we must utilise when creating colour schemes.

I don't think we make enough of how display-adapted Spinosaurus aegyptiacus was, and how weird that is for not only a giant dinosaur predator, but any giant predator. Here, a gaggle of Spinosaurus show off their sails and tails, display structures (well, probable display structures, in the case of the tail) that use almost every inch of their axial length for showing off. So how does this fit into your "big theropods were camouflage-colour dominant" narrative, smart guy?

The elephant in the room here, of course, is Spinosaurus, which is highly unusual for being a giant apex predator with the same tailor as a peacock. This was a carnivore with an unprecedented disregard for remaining inconspicuous or having an anonymous body profile. For all the controversy over this animal, one aspect we all agree on is that its enormous sail was a sociosexual display device (see Hone and Holtz 2021 for references and discussion). Doesn’t this doesn’t torpedo the wider point being made here about predator size and possible camouflage needs? On the contrary, it might support it. As something straddling the terrestrial-aquatic realm, normal rules about camouflage and crypsis may not have applied to Spinosaurus. We see this evidenced in modern times in that the "rules" of camouflage in terrestrial settings are not the same as those of aquatic habitats (Caro 2013), and we should probably allow for, or even expect, some weirdness from animals operating at that interface. The atypical ecology of spinosaurids may have liberated them from the adaptive pressures experienced by purely terrestrial dinosaur predators, allowing them to become more ornamental and spectacular. Perhaps the fishy prey of Spinosaurus barely saw the full outline of their largest predator, an especially viable idea in the (I think, superior) “giant heron” ecological model favoured by several authors (e.g. Hone and Holtz 2021; Sereno et al. 2022).

Pigment availability

Moving on, can we get a sense of the skin pigmentation available to giant theropods, thus letting us know which paints/colouring pencils/digital palettes to crack open? Here, we have to think about the availability of environmental pigments, like carotenoids. Many readers will know that animals cannot create all the pigments used in their integument and that some are obtained through eating plants or microbes. Carotenoids and other environmental pigments create some of the most vivid colours seen on animals today, including hot reds, bright oranges and canary yellows. But environmental pigments are hard to source in terrestrial settings, to the extent that even tiny songbirds compete with one another to source them (Blount 2004; Biard et al. 2005). Outside of specialist ecologies, the most famous being that of flamingoes, larger terrestrial animals tend to make do with pigments they can manufacture themselves, such as melanin. This is one reason why so many terrestrial animals are earthy tones, such as greys, blacks, browns, orange-reds, and white (where pigment is withheld). But structural colour, features of skin, scales and feathers that manipulate light to create colour without pigmentation, has also been developed across all vertebrates and is exploited to produce greens and blues. In all probability, it’s from these basic pigment and structural palettes that giant theropods were deriving their hues. Unless conditions of the past were very different to those of today, it’s hard to imagine multi-tonne terrestrial animals finding enough carotenoids to develop large patches of particularly intense pigmentation.

Specifics of patterning

Our discussion raises a notch in complexity as we move to consider giant theropod skin patterns, even if we stick within the camouflage-dominant framework outlined above. Concealment strategies are adapted to specific habitats, predation styles and prey types because no one system is universally effective. Indeed, one of the few constant rules of camouflage — that, no matter how perfectly a crypsis strategy works on a stationary animal, movement always gives the game away (Pembury Smith and Ruxton 2020) — is of little use to us here because we don’t know where and how big theropods hunted. The concealment strategy of an endurance predator, one that simply hounds its prey tirelessly, waiting for it to become vulnerable from exhaustion, might be different to that of an ambush predator that relies on surprise, springing at its prey at the last moment for a short chase.

These are only the first factors to consider. Predator colours are also modified by the time of day the predator tends to operate, as well as their position in the food chain: some have to be worried about being prey items themselves. And that, in turn, is altered by the colour schemes that can be created by different integument types (e.g. fibres vs. naked skin vs. scales), as well as the functional impacts of pigmentation. Darker pigments, for instance, can protect skin from harmful UV rays and may have antibacterial properties but, conversely, also absorb more solar heat and increase an animal’s thermal load (Walsberg 1983; Caro 2005; Caro and Mallarino 2020). There’s a lot to think about here, and the fact we still can’t account for these and other variables reliably in living animals is why biologists still consider our knowledge of animal colour to be fairly limited. It goes without saying that, if we’re still working out what’s happening among living species, robust predictions of camouflage patterning in extinct animals are way off.

The colour schemes we give our dinosaurs have functional implications beyond interacting with other animals. This dark, adult Tyrannosaurus would be well-protected from solar radiation by its dark skin, but it would absorb a lot of heat in direct sunlight.

Nevertheless, we may be able to narrow down some possibilities for giant theropods by looking at what works for large modern predators. Most employ background matching, where their skin tone approximates that of their surroundings, or else they use countershading, where dark upper regions and lighter undersides disrupt the formation of shadows, diminishing contrast with the background (note that this is disputed by some, there is actually a fair amount of controversy around countershading function: see Ruxton et al. 2004; Rowland 2009). Other predators use disruptive colouration, where high-contrast colours break up body outlines and disguise distinctive features such as eyes. Unlikely strategies for big theropods are masquerading tactics: attempts to match unexciting objects like rocks or twigs. To pull off this illusion, masqueraders have to resemble something of equivalent size and shape, and that becomes harder at larger sizes, perhaps explaining the absence of this method among large terrestrial predators today. This strategy is distinct from mimicry, where an organism adopts the appearance of another species to be misleading about its true nature (Skelhorn et al. 2010).

With several patterning options on the table, progressing further with this discussion is only possible if we start making assumptions about giant theropod ecology, pushing us further into the realm of inference and speculation. But we can ground ourselves by considering the results of studies into camouflage function and performance. For instance, if countershading does indeed work to disrupt shadowing, then studies show that a sharp, high-body colour transition would work better in an open setting than a more gradual colour change lower on the flank, which obscures animals more effectively in forested settings (Vinter et al. 2016). We generally see more uniform, low-contrast colours on big animals in open habitats because large patches of colour generally don’t conceal animals as well in woodlands (Pembury Smith and Ruxton 2020, although flat-grey elephants are reportedly remarkably difficult to find once they enter forests — see Caro 2013). Conversely, high-contrast patterns seem to work better at hiding animals in vegetated or otherwise busier environments.

Baby tyrannosaurs, barely a metre long, with cryptic colours that help them blend into the forest floor. In all likelihood, baby tyrants were at high predation risk and it seems reasonable to assume they used camouflage tactics to avoid being eaten. But the colour schemes of infants may not have worked so well for their gigantic parents, nor even older juveniles or subadults. Might tyrannosaurs and other giant theropods have tracked through multiple colour morphs en route to somatic maturity?

We can consider things like the age of our restoration subjects, too. In scaly animals (the only skin type we currently have direct evidence for in giant predatory theropods, even if we can’t rule out the possibility of some protofeathering), colour vividness tends to reduce with age (Olsson et al. 2013). This change may not just be physiological, but also adaptive. The juveniles of all animals, including apex predators, are targetted by carnivores and their colouration has to be multi-functional, hiding them from predators as well as — in precocial species — their prey. This is often achieved with disruptive patterning. Stripes, spots, bars and other features may serve an additional role, achieving a “motion dazzle” effect that confuses predators about animal speed and direction, or draws focus to less critical anatomies, like tail tips (Murali & Kodandaramaiah 2016). Dazzling capacity diminishes at lower speeds and agility, and is thus less useful in larger animals (Pembury Smith and Roxton 2020), perhaps partly contributing to the dulling of living reptiles as they approach adulthood. We should not imagine that juvenile theropods transitioned to their adult colours straight away, however. It took decades to grow gigantic theropodan predators and, in all probability, the route to adulthood was via several different ecological niches (e.g. Holtz 2021), each of which may have had different adaptive pressures on colouration. So maybe giant theropods had several colour schemes throughout their lives, and we should render them as being colour-adapted to their various age-specific lifestyles? We could go on listing the adaptive aspects of different animal skin patterns all day, but you get the idea. There's a lot of camouflage science we could factor into our reconstructions, even if we can't ever know the real colours and patterns of our subject species.

So... does animal colour science help us in palaeoart?

Background-matching, age-dulled Tyrannosaurus rex takes on a countershaded, partly disruptively-coloured Edmontosaurus annectens. These guys are mainly here to stomp about and wake people up with some Hardcore Dino Action™ in case anyone has drifted off when reading this long, long post.

Let’s conclude by returning to our main question: can studies of living animals constrain our speculations about the colours of dinosaurs, or will colour restorations forever remain a crapshoot when we don’t have palaeocolour data? Here, we've extrapolated the findings of predator-specific colour studies to giant, terrestrially-hunting theropods and, based on these, we've suggested that large dinosaur predators...

• were likely under very strong pressures for crypsis
• probably didn't load their skin with many environmental pigments
• likely expressed background matching, countershading or disruptive patterning, depending on their specific ecologies
• may have had several colour schemes throughout their lives as their ecology changed with age

Down the line, we can discuss the merits of these predictions — do remember that you're reading a secondary take on all the science discussed here and that you may come to different interpretations based on your own literature crawl. But it's not these specific findings that are most important here. Rather, it's that this case study shows what animal colour science can offer to the process of restoring one type of extinct animal, as well as its broader potential for focusing our loosely-constrained applications of colour within palaeoart. The points made above, or others like them, do not give a colour scheme for giant theropods, but they do suggest that some concepts are more likely than others, and even rough guidance isn't to be sniffed at when we're otherwise running virtually blind. It's strange that palaeoartists are often able to point to core palaeontological studies for interpreting fossils, and core anatomical studies for depicting anatomy, but we don't generally talk about or know the same literature on animal colour. I wonder if it'll eventually be worth keeping up with developments in this field as much as we do new fossil and anatomical data — if we’re not at this point already. The result can only be more scientifically credible and realistic artwork, and that's a win for everyone.

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References

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Palaeoartist interview: Johan Egerkrans

Palaeoart has never been a particularly diverse artform. Since the early 1800s most palaeoartists have pursued art attempting to depict fossil animals in realistic ways, with stylistic variation mostly along the spectrum of how obvious our brush strokes and pencil lines are, and how much detail we add. In recent decades we've seen artists deepening their dedication to realism with hyperrealist palaeoart, artworks which look like they've been snapped by high-speed cameras with crisp focuses and ultra-high levels of detail.

But not all palaeoartists are taking this approach. Some take a step away from not only high levels of detail but also realism, producing palaeoart with a more stylised and even abstract bent. Though few in number, the growing roster of ‘stylised’ palaeoartists represent an exciting new frontier for palaeoart. In varying artworks along spectra other than tidiness and detailing, these artists are producing unconventional works recalling pop art, classic western animations, heraldic crests, perspectiveless Medieval art and more. Among the most fascinating aspects of these works is their capacity to maintain respect for scientific credibility even when producing stylised, non-realist art. The forms may be simple or sharply angular, the colours may be garish, but we can still tell what the subjects are, what they are doing, and get a sense of their anatomy.

...which brings us to Johan Egerkrans's Alla tiders dinosaurier. If you like stylised palaeoart, you should check out this book. 

Swedish artist Johan Egerkrans is part of this emerging group of unconventional palaeoartists. Emerging onto the online palaeoart scene only recently, his work has already generated a fanbase and widespread acclaim. It's easy to see the appeal of his creations. Distinctively angular, full of personality and recalling great works of American animation, his digital artworks emphasise and almost caricature the form of fossil animals without undue distortion of their form or disregarding fossil data. Attention to details, anatomy and colours make his work interesting to look at despite it's simplicity compared to traditional modern palaeoart. We're not just seeing generic cartoons of fossil animals, but highly stylised versions of contemporary, scientifically credible palaeoart, informed by a clear appreciation for modern wildlife and the natural world. Notice the pupil colour change between his adult and juvenile Microraptor (below), variable integuments on Gorgosaurus (above), fine attention to animal poses and behaviour, and so on. His use of traditional compositions and poses prevent his work becoming overbearing: in this regard, his work is less intrusive, and even perhaps less cartoony, than some artists employing ‘realistic’ animals in hyper-dynamic poses and compositions.

Egerkrans' parent and offspring Microraptor. Look past the stylisation and this is a pretty accurate take on Microraptor anatomy, right down to the iridescent black plumage. Note the pin feathers and dark pupil on the juvenile - very sensible speculations for juvenile maniraptorans. © Johan Egerkrans.

Each Egerkrans work radiates personality: his animals have real character, and it’s almost impossible not to imagine them taking part in animated vignettes. Several of his works have a strong sense of mischief and dark humour, another rarity among palaeoartworks. I’m particularly tickled by his scene of a capybara running away from terror bird Titanis (below): the bird has a mania that captures real birds at their most frantic and chaotic, while the drab mammal looks overwrought, panicked, but also like it’s going to write a strongly worded letter to the Daily Mail about all this. Comparisons of Egerkrans’ creations to stylised fossil animals rendered for the big screen are inevitable, and mostly leave us wondering what the heck everyone else is doing wrong. Hollywood, give this man a job!

Titanis and capybara star in Hilarious Scene of Violence. Capybara won an Oscar for its eyebrows. © Johan Egerkrans.

Johan was kind enough to send me a copy of his recent book, Alla tiders dinosaurier, which I thoroughly recommend you check out. There’s no English translation at the moment (one might happen at some point) but the artwork speaks volumes alone and the design and print quality is excellent - it's a nice book to have, even if you're unable to read the text. The follow up, Flygödlor och havsmonster, which focuses on marine reptiles and pterosaurs, is due out later this year. Both are published by B Wahlstroms, and can be purchased from Bokus and other Swedish book retailers (sorry, American readers, there are complications around shipping these books to the USA at the moment). You can check out the art of both books on Facebook, Artstation and Johan's blog. If you're Stockholm-based, you can also check out a dinosaur exhibition featuring the Alla tiders dinosaurier work, which is running until the end of September.

Earlier this month I asked Johan if he’d like to chat to me about his art, books and palaeoart philosophy, and he’s taken time out of his schedule to give the following interview. With thanks to him for taking time to respond to my questions, it’s time for me to stop gushing about his work and hand you over to the man himself…

MW. You’re quite new to the palaeoart scene, but have landed an instant fanbase with your highly distinctive artwork. Can you give us some insight into your artistic background and what brought you into restoring dinosaurs, pterosaurs and so on?

JE. Hi Mark! Thanks for having me on the show!

I started out as a concept artist and, like most people in that field it seems, I´ve nursed  a deeply rooted fascination for paleoart since... Well, forever I guess. At the age of four my dad gave me Burian´s seminal art book “Life Before Man” and that was it; I was hooked and filled countless A4 sheets with scribblings of dinosaurs, therapsids, pterosaurs and other extinct beasties. I´ve still got that same cherished tome in my bookshelf, worn and coming apart at the seams.

Fast forward to the early 2000´s when I got my first fulltime job as an illustrator concepting for a small computer game outfit called Idol here in my hometown Stockholm. There I did designs for monsters, robots, spaceships and stuff like that. A high point was when I got to draw a series of - listen to this - demonically possessed cyborg dinosaurs!  That´s about as awesomebro as things can get. Take that Michael Bay!

I was always had a talent for mimicking different art styles, which came in very handy at that job - one month you did a superhero game in a highly stylised Bruce Timm style, another month it was horror inspired by Clive Barker, Frazettaesque fantasy or something completely different. I really got to flex those versatility muscles in that environment.

Anyway, after a couple of years Idol went belly up, as small computer game outfits are wont to do. I became a freelance illustrator and found myself working more and more with children´s books. In 2013 Nordiska väsen/Vaesen was released - a book about creatures from Scandinavian folklore that I wrote and illustrated. That really was a watershed moment, as the book did rather well (still does - it's sold over 40.000 copies in Sweden alone so far). After that success I had a certain amount of freedom and one of the things I wanted to do was to go back to my paleoart roots in some fashion. The first such project was a children´s picture book called My first book of dinosaurs. It was originally intended to be a rather tongue-in-cheek affair and the initial pictures were intentionally tropey (large theropod roaring on cliff, cassowary Oviraptor). I did take care to stay off the beaten path though so, unusually for a book aimed at young children, there wasn't a T. rex or Triceratops in sight - I went with Giganotosaurus and Styracosaurus instead.

Mention the tropes, and they shall appear. Egerkrans' Smilodon bellowing off a cliff (or maybe suffering a major case of lockjaw). It's difficult not to see this as satirising the most traditional means of restoring sabre-toothed cats: the lower jaw stretched so far as to make its tissues near invisible, and the skull arcing upwards to attain more ferociousness. Image © Johan Egerkrans.

Pretty soon my science geek side kicked in - I did more and more research and realised I wanted the reconstructions to have a certain amount of scientific accuracy, even if the book was aimed at toddlers. The cartoony stylised style I had chosen for the book could be tweaked into some something more “serious” while still retaining the whimsy and charm of those first illustrations. My first book of dinosaurs was followed by a another one about Cenozoic beasts and by this time I had gotten wind of the All Yesterdays movement and had started following a bunch of paleoblogs (this one and Tet Zoo among them). This new wave of paleoart and the philosophy behind it appealed to me. My editor and I decided to do a “real” pop science book about dinosaurs which was released as Alla tiders dinosaurier ("Dinosaurs of All Ages") earlier this year. I´m currently racing towards the finish on the follow up about pterosaurs and Mesozoic marine reptiles.

MW. Strongly stylised palaeoart is rare, perhaps because we focus so rigidly on precision and scientific credibility in our reconstructions. Where do you draw the line between style and adherence to science, and are there cases where you’ve thought ‘screw science, this looks cooler!’

JE. My aim, in a way, is to do what Disney animators did in films like The Jungle Book or The Lion King. Now, Shere Khaan might not be realistic per se, but the design is informed by a deep understanding of tiger anatomy, and what tigers are like - their “essence” if you will, with the risk of sounding a tad pretentious. Thus Shere Khaan becomes the tigeriest tiger around as far as I´m concerned. My paleoart sort of tries to do something similar - only with extinct animals (though I´m nowhere near as talented as those old school Disney animators). To capture that “essence” you sometimes got to break the rules a bit. It´s a “know the rules to break the rules” kinda deal.

It´s a bit like caricatures come to think of it. People often find it easier to recognise a celebrity from a (well made) caricature than from a photo because the drawing exaggerates that person's distinguishing features. In a similar way stylisation allows me to focus on what’s distinctive about a certain species/genus and bring that up to front.

Parvicursor, from Alla tiders Dinosaurier, is a great example of Egerkrans' capacity to find the essential elements of form in an extinct animal and project them through a strong visual style. © Johan Egerkrans.

Another advantage is that it allows me to remain vague when we’re uncertain about some feature of an animal's anatomy. Take for instance the recent dispute whether tyrannosaurs had lips or croclike exposed teeth. The simplified style allows me to draw something in-between, should I so wish, and leave it open to interpretation. That doesn’t mean I do this all the time and never takes a stand, but it remains an option.

A lot of paleoart seems rather overworked. I´m hardly the first to voice this opinion but meticulously rendering thousands of  tiny scales in a dinosaur picture doesn't necessarily make said picture more accurate. Sometimes it´s the complete opposite where hyperrealism only serves to create the illusion of scientific accuracy. I tend to prefer sketchier, looser paleoart - by artists like John Conway, Simon Stålenhag and of course Zdeněk Burian - where the emphasis lies on movement, mood and communicating that aforementioned essence of an animal - what it felt like.

My most common “screw-you-science” is probably the eyes. The peepers of my stem-birds are more mobile than they probably were in real life; they move around and look at things in a human, or at least mammalian way. Avian eyes are usually fixed in a perpetual stare which makes them come off as either vexed or insane (or both). That might be precisely what you’re after, but often you’re looking for something different. I almost always give the animals discernible pupils as we humans are geared to interpret that as more affective than-all black eyes. Windows to the soul and all that.

MW. Your reconstructions are full of personality and humour. I find it very easy to project emotion onto your subjects. Is this something you deliberately seek with your work? Do you render each image with an idea about what each animal is thinking?

JE. I´ve always had a flair for characterisation. It just sort of happens no matter what I draw, be it a robot, a dragon or a lone animal hanging about doing nothing. They always end up seeming to be up to something (my subjects often look rather smug for some reason, apparently it´s my go-to emotion). There´s a hint of anthropomorphism but I try not to overdo it. It´s just little things like an eye ridge tweaked to look as if the animal is raising it´s eyebrows or the hint of a smirk at the corner of the mouth. It should only be just enough to help the viewer empathise with the subject.

MW. The colour choices of your artwork are interesting, blending ‘realistic’ animal colour schemes with background hues rarely seen in palaeoart. It works very effectively, creating a strong sense of atmosphere. Can you take us through your approach to choosing animal colouration and blending these with often contrasting backgrounds?

JE. I always start with the animal itself and let their colouration dictate the tones of the background. The aim is to give them striking, simple colour schemes that still comes off as believable. Once the animal is painted I start with the surrounding environment, which on the whole is a rather intuitive and organic process. I play around in Photoshop until I land in something that works.

The colour choices and compositions are highly influenced by animation backgrounds, especially in the way the scenes are framed. There´s a lot of colour theory at work as well - complementary colors (often good old orange and teal) or split complementary colours (like red and blue) in different overlay layers make the animals “pop” from the background. A cool coloured animal will be framed by a warmer environment and vice versa.

Dimorphodon meets a neighbour (notice the keratin crest on the lower jaw of Dimorphodon - most artists miss that). In addition to showing the personality common to Egerkrans' work, this piece also shows the mix of realistic animal colouration with striking, pseudorealistic background colours. In fully realistic art, this might not work, but here, it does. © Johan Egerkrans.

MW. To me, your palaeoartworks recall some of William Stout’s illustrations. Both have a distinctive, non-realist style, interesting colour schemes and emphasis on the animal subjects. Is Stout an influence on your work?

JE. Very much so. I've always loved his work and his approach to paleoart. His creatures have tons of character and the draughtsmanship is sublime. They’re admittedly a bit skeletal at times but they make that up with personality. That I’m partial to Stout is hardly a surprise, as we're both inspired by the same old masters. Even if it's not obvious in my paleoart, a lot of my work takes cues from turn of the century illustrators like Arthur Rackham, Dulac and John Bauer, just like Stout's art.

MW. The work you produce is included in educational books. How do you think style impacts the scientific or educational prospects for palaeoart?

JE. The illustrations are not intended to be photoreal and that´s sort of the point. It´s obvious that they're an interpretation which forces the viewers to do part of the reconstruction in their own heads. That hopefully gets their imagination going which is the ultimate goal - to connect and get people interested. To make science fun.

The chosen style also saves me from meticulously rendering those thousands of tiny scales and retain my sanity, so that´s a huge plus.

MW. Do you ever stray from your signature style? Will we ever see a ‘realistic’ Egerkransian dinosaur?

JE. As I´ve mentioned before I always adapt my technique to the project at hand and this is just one of several styles I utilise. It´d be interesting to do a paleoart project in a more realistic vein, though I think there´ll always be a certain amount of stylisation. I´m not a realist painter and never will be - others have got that down already.

Umoonasaurus and chums. The barnacled fallen trees turns this image from just another Mesozoic marine scene into something much more atmospheric. © Johan Egerkrans.

MW. I’ve seen that you get a lot of scientific feedback on Facebook posts, a source that many palaeoartists – professional and amateur – can be wary of because of misinformation and confrontational internet users. How useful do you find social media to shape your art, and have you encountered much hostility?

JE. I was flabbergasted at how overwhelmingly positive the response was when I posted my first drawings on the Facebooks. Especially from the academic community. There´s been very little hostile or dismissive remarks - in general people seem to take the works seriously, as ‘proper’ paleoart.

The feedback is often extremely helpful - there´s lots of very well informed academics hanging about (you yourself and Darren Naish to mention just a few) and you quickly learn to sift the good advice from the bad or opinionated. I approach the forums as a sort of quick and dirty peer review; I´m not an expert and get things wrong all the time and if there´s something wonky someone is bound to point it out. As the ambition is to be as accurate as possible, within the limitations of the style, I try to surround myself with people who actually truly knows about this stuff. As luck would have it a lot of people I admire have proven to be more than willing to help out with comments, constructive criticism, links to papers and by just being supportive in general.

MW. When are you going to get Hollywood on the phone to make your work into a movie? They already look like they’re stills from some epic animated film about Mesozoic life. And they owe us, frankly, after The Good Dinosaur.

JE. I´m still waiting for them to get the straws out of their noses and give me a call. Bastards.

Guanlong and some sort of impudent Mesozoic mammal. Note how the Guanlong is strikingly and variably coloured, and yet still looks grounded. Bringing bright colours into the Mesozoic doesn't necessarily mean painting entire animals in lurid shades. © Johan Egerkrans.

MW. Finally, where’s the best place to find your art and support your work? And how long do we have to wait until your next book?

JE. You can follow my public facebook account “Johan Egerkrans - Illustrator” where I post about new projects and upcoming events like signings. Then of course there is the Paleoartists Facebook group where I´m pretty active.

I´ve also got a blog at http://johan-egerkrans.blogspot.se/ and an Artstation page https://www.artstation.com/artist/egerkrans.

My books can be bought from www.bokus.com or any other Swedish book retailer. You should be able to order them from there if you live in Europe but it's trickier in the States due to the fickle nature of the U.S. customs. Hopefully Alla tiders Dinosaurier will get an English edition at some point, but nothing's set at the moment.

The next book Flygödlor och havsmonster, about your favourites the pterosaurs (and their marine contemporaries), will be out in Sweden this fall. At some point I´d very much like to do a book about Permian and Mesozoic stem mammals (gorgonopsids are hands down my favourite prehistoric animals), but sadly it is a rather tough sell…  

MW. Johan Egerkrans, thanks very much!

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