Controversial ceratopsids revisited: woolly Pachyrhinosaurus and scavenging Styracosaurus

Spurned on by a print request, I've spent free time this week revising two images of ceratopsids which may be familiar to long-term readers: my woolly Pachyrhinosaurus perotorum and scavenging Styracosaurus albertensis. The former is now just over two years old, and the latter a whopping eight years old - wow, have I really been messing about with internet palaeoart for that long?

Maastrichtian Alaska was quite chilly, but woolly Pachyrhinosaurus perotorum doesn't care. See this post for the original image and exploration of the concept shown here. Prints are available.

Because I appreciate some folks are fond of my original paintings, I haven't deviated too far from the original compositions and instead just added more detail, tweaked colour values and tidied up some sketchy areas. I'm very conscious of not 'pulling a Lucas' on my old work. Most importantly, the science has been improved/corrected: the cranial morphology of Pachyrhinosaurus perotorum is now correct to that species (like a doofus, I based the morphology in the original image on a different Pachyrhinosaurus species) and the tyrannosaur in the scavenging scene is appropriately filamentous. Thanks to Darren Naish, Zachary Miller and Christian Kammerer for discussions of Styracosaurus horn shape.

Have these depictions have been supported or refuted by any new discoveries? As far as I'm aware, skin impressions still remain elusive for Pachyrhinosaurus, although new data has emerged on the facial integument of juveniles (Fiorillo and Tykoski 2013). The 2014 discovery of Kulinadromeus and its assortment of filament-like scales, true filaments and other integumentary oddities (Godefroit et al. 2014) might indirectly add credence to the idea of shaggy ceratopsids, however. Along with Psittacosaurus and Tianyulong, Kulindadromeus shows that the evolution of ornithischian integument was complex, that single animals can bear a suite of different integument types, and that the assumption of dinosaur skin being ancestrally scaly is uncertain. The weird scales in (unpublished) skin impressions of Triceratops are further evidence that 'one skin fits all' approaches to reconstructing these animals are likely flawed, and that even clades with relatively limited anatomical disparity - like ceratopsids - had diverse integuments. Thus, the idea that some members of Dinosauria may have looked very different to our traditional interpretations is being strengthened by genuine data, and shaggy arctic ceratopsids remain a fun extrapolation of that concept. For further discussion on these points, check out my discussion of version one of the woollysaur painting.

The Campanian centrosaurine Styracosaurus albertensis scavenges the remains of a tyrannosaurid. He was going for warpaint on his face, but he ended up at 'Tonto'. For fun, the original 2007 version can be seen here. Prints are available.

What of scavenging ceratopsids, as in the reworked 2007 image of a tyrannosaurid-eating Styracosaurus? Ceratopsid omnivory has yet to be explored in the technical literature and, to my knowledge, remains best represented by a short paragraph in Paul (1991). More recently, Mallon and Anderson (2013) provided reasoning for why ceratopsids were not predatory animals, although their discussion seems to consider 'carnivory' synonymous with 'predation': opportunistic scavenging or omnivory are not explored. This leaves most discussion of ceratopsid scavenging online, and several famous denizens of the online palaeontological community seem to support it. Back in 2007 I wrote a long essay substantiating the idea. That essay is no longer online*, but the argument is pretty straightforward:

*After eight years, I figured it's time to archive my old Flickr stream. The bulk of the content there is not representative of modern science or a good representation of my work, so it's been taken offline. I won't pretend I'm not a bit sad to do so, but there's obviously reason for bringing internet searches to my best, most recent work, not images I created when first learning how to paint.

1. As is well-known, a number of modern herbivores eat animal remains on occasion. This may reflect nutrient stress (thought to explain carrion use by hippos, which is not as common as 'common knowledge' might suggest) or else a method of supplementing a mineral-deficient diet (as in deer, cows, giraffes and a host of other hoofed mammals - Hutson et al. 2013). Remarkably, some cases of hippo carnivory involve the hippos killing animals first, and they will also scare other carnivores from kills to obtain carcass access (Dudley 1998). Of further interest is that entire herds of hippos will chew on carcasses when available - these are not the acts of rogue, aggressive or aberrant individuals (Dudley 1998). Note that studies on the carnivorous tendancies of generally herbivorous animals are in their infancy, and it may be that this behaviour is more common and opportunistic than we currently realise.
2. Other species, such as pigs, ingest animal matter as part of their normal diets. Studies on some pigs suggest 28% of their diet is derived from animals, either being invertebrates or carrion (e.g. Thomson, and Challies 1988). There is no reason to think that large extinct animals were incapable of comparable omnivory, but we restrict most discussion of it to smaller dinosaurs and pterosaurs. We can predict that such animals should have jaws mostly adapted for herbivory (e.g. teeth suited to browsing and grazing, long 'cheek' toothrows, vertically displaced jaw joints etc.) but would also have some means to process animal remains (e.g. crushing teeth to break bones, caniform teeth or sharp beaks for ripping meat etc.).
3. Ceratopsid jaws certainly belonged to primarily herbivorous species capable of chewing their food, but their approach to herbivory was unusual. Their teeth and jaws, unlike other herbivorous dinosaurs and mammals, were incapable of grinding plant matter. Instead, they sliced food into pieces, their teeth sliding vertically past one other like scissors. Ceratopsid beaks are also unusually deep and narrow compared to other dinosaurian herbivores, and recall the beaks of parrots in many respects. The beaks of these birds are famously powerful, enabling their owners to access a range of nuts, seeds and animal matter (e.g. Greene 1999). The diet of of ceratopsids has been questioned by palaeontologists because chopping plant matter is not common among modern herbivores. To the contrary, most food slicers are carnivores - meat is easier to chop and slice into easily digested chunks than it is to grind into a paste. One sensible suggestion is that ceratopsids ingested particularly fibrous, woody plant matter (see Mallon and Anderson 2013 and references therein). We might imagine them devastating Cretaceous shrubs, removing entire chunks of tree - leaves, branches and bark - with each bite, or overturning plants with their huge heads to access their roots and tubers. However, it is odd that their jaws aren't more convergent with those of other herbivores, as grinding mechanisms have developed so many times in multiple tetrapod lineage and might be considered optimal for breaking down plant matter. So, maybe ceratopsid jaws were used for more than simply eating plants, and their shearing teeth and hooked beaks are the traits of omnivory we mentioned above, equally capable of slicing plants and animal remains. Opening carcasses, snapping smaller bones and slicing meat was almost certainly possible with their jaws and beaks, and we might imagine ceratopids as Mesozoic variants of pigs: largely herbivorous species with opportunistic carnivorous tendencies, and certainly capable of competing with strict carnivores for carcass access. The possibility that they could occasionally kill other animals for food, as demonstrated by the aforementioned hippos, is not unreasonable.

Back in 2007 I mentioned a possible smoking gun for this idea - a rumoured Psittacosaurus specimen with bony gut content. Since then, it's become apparent that that specimen either doesn't exist, has disappeared or has otherwise been forgotten about - it's best to consider that an unsubstantiated rumour for now. Despite this, I still think the concept of ceratopsian omnivory has legs: maybe a technical paper on the topic would be worthwhile.

Bumblebee Conservation Trust charity prints: an update

In my last post I mentioned you can buy a print of my Tyrannosaurus vs. bees painting and donate money to the Bumblebee Conservation Trust. I'm happy to say £55 has been raised in the last week for this cause, and thanks to those who've bought in. It would be great to make even more money however: if you'd like to contribute, find out more here.

Of course, prints are available for all my other work too, including the ceratopsid pieces above. Contact me at wittonprints@gmail.com to order one, and check out this page for prices and other details.

References

• Dudley, J. P. (1998). Reports of carnivory by the common hippo Hippopotamus amphibius: short communication. South African Journal of Wildlife Research, 28(2), 58-59.
• Fiorillo, A. R., & Tykoski, R. S. (2013). An immature Pachyrhinosaurus perotorum (Dinosauria: Ceratopsidae) nasal reveals unexpected complexity of craniofacial ontogeny and integument in Pachyrhinosaurus. PloS one, 8(6), e65802.
• Godefroit, P., Sinitsa, S. M., Dhouailly, D., Bolotsky, Y. L., Sizov, A. V., McNamara, M. E. & Spagna, P. (2014). A Jurassic ornithischian dinosaur from Siberia with both feathers and scales. Science, 345(6195), 451-455.
• Greene, T. C. (1995). Aspects of the ecology of Antipodes Island Parakeet (Cyanoramphus unicolor) and Reischek's Parakeet (C. novaezelandiae hochstetten) on Antipodes Island, October-November 1995. Notornis 46: 301-31
• Hutson, J. M., Burke, C. C., & Haynes, G. (2013). Osteophagia and bone modifications by giraffe and other large ungulates. Journal of Archaeological Science, 40(12), 4139-4149.
• Mallon, J. C., & Anderson, J. S. (2014). The functional and palaeoecological implications of tooth morphology and wear for the megaherbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta, Canada. PloS one, 9(6), e98605.
• Paul, G.S. (1991). The many myths, some old, some new, of dinosaurology. Modern Geology, 16: 69-99
• Thomson, C., & Challies, C. N. (1988). Diet of feral pigs in the podocarp-tawa forests of the Urewera Ranges. New Zealand journal of ecology, 11, 73-78.

Deconstructing All Yesterdays, or How palaeoart is flawed, but everything's cool

It's time to face facts. Try as we might, we will never reconstruct long extinct animals accurately. We may be able to cobble together fairly accurate images of Pleistocene mammals through analysis of their frozen remains and heavy reliance on closely related modern species, but the appearances of species extinct for millions of years are beyond are grasp. Our problems are far greater than the most common complaint, that we simply do not know what colours they were. So much data on soft tissue distribution, muscle bulk and integumentary structures are lost through the death, taphonomic processes, fossilisation and exhumation of extinct creatures that very little can be said for certain about their actual life appearance. The remains that weather the fossilisation process - skeletons and shells - only provide the bare minimum of information about the life appearance of their owners, and it seems that osteological correlates - features of skeletons that betray the presence or development of certain soft-tissue structures - are often ambiguous or unreliable. And we haven't even mentioned the problems with trying to deduce behaviour from bones alone. (Image above: the underlying sketch to this controversial painting of a carrion-eating, bristly Styracosaurus. From 2007.)

This is one of the messages I've taken from the presentations, internet articles and new book All Yesterdays. The brainchild of John Conway, C. M. Kosemen and Darren Naish, this minor internet phenomenon needs little introduction to most of the readers here (if you're unfamiliar with it, check out the provided links for background info). This opinion contrasts with the attitudes of some palaeontologists and artists, who consider our abilities to reconstruct extinct animals fairly decent and reliable. Pains me as it does to say it, but I have to agree with the All Yesterdays chaps. There's simply too much anatomical and behavioural detail lost to time, and we're never going to get that back.

The All Yesterdays project seems to be the result of looking into this deep abyss of lost palaeontological data. But rather than staying safe at the edge, Conway et al. have dived in, exploring the virtually infinite possibilities of ancient animal reconstructions, critiquing the rationale and methodologies of palaeoart and questioning its very purpose. The results are novel, highly creative, insightful and thought provoking, and should be given serious thought by anyone interested in the palaeobiology and depiction of extinct animals. In short, All Yesterdays argues that modern palaeoart fails one of the only tests we can apply to it, that many depictions of extinct animals compare poorly against the morphological and behavioural diversity of modern species. Specifically, modern palaeoart is too conservative, frequently depicting set behavioural patterns for some species (e.g. the Tenontosaurus vs. Deinonychus meme) and 'shrink wrapping' skin over the skeletomuscular system without any consideration of other soft-tissues. All Yesterdays argues that this can be rectified, in part, through bold transference of modern animal anatomy and behaviour to extinct species, which fills the gulfs of missing data and creates more plausible concepts of life in the past than by following the current, perhaps overly conservative palaeoart methods. The word 'concept' is important here, as this is all we can hope to realistically hope to achieve in palaeoart. For all our efforts, our reconstructions will probably never depict these animals exactly as they were in life, and we just have to live with that. Science will hone and refine our concepts to more closely resemble what was once reality, but most of the details we need to exactly reconstruct ancient worlds are unlikely to ever emerge. There will always be several plausible ideas for the life appearance of extinct creatures*, and we should focus on exploring these concepts to reproduce believable renditions of animals, not ignoring because of (probably) unobtainable data or because they stray from established ideas.

*Is there more than one way to reconstruct a fossil animal? According to some, no, but others would disagree. I think the answer lies somewhere inbetween. The skeletomuscular system of extinct animals may be reconstructed more-or-less correctly from fossils of some species but, as we'll see below, this is only half of the story.

But we must be careful with this liberation of creativity in a science-based discipline. All Yesterdays calls not for recklessness in palaeoart, but confidence, allowing extinct animals to be diverse and unusual, but still constrained by what is known for their evolutionary history and anatomy. The payoff for this confidence is that the All Yesterdays project often portrays extinct animals in a more convincing and realistic manner than much of the work we're familiar with. For example, John's tripodal Therizinosaurus is just as plausible, scientifically speaking, as a more traditional version, but is 100 times more believable. John's Camarasaurus rolling in mud is just as plausible as the hundreds of illustrations of this animal standing and eating, and Memo's super-stocky Lambeosaurus is entirely consistent with fossils of this species. The results are just as valid as hypotheses of appearance and behaviour - arguably moreso - than the ultra-conservative reconstructions currently dominating palaeoart. Erring on the side of caution is still an error, and some of the entrenched, 'conservative' reconstructions of ancient life are actually harder to substantiate than the seemingly bolder ones.

Elements of what I'll call 'the All Yesterdays philosophy' have been creeping into palaeoart for years, but I think Conway et al. have burst the dam here, highlighting the need for a fresh approach in how we approach the reconstruction of extinct life. So far as I see it, there are four points to consider in the All Yesterdays philosophy that both palaeontologists and palaeoartists need to embrace: when to apply it, the composition of our images, what it means for animal appearance, and its application for extinct animal behaviour.

Application
All Yesterdays is entirely about reconstructing long extinct animals with no ecologically similar modern relatives. We don't need to extrapolate data wildly for relatively recently extinct species with lots of closely related modern relatives, and doing so will probably make our work less accurate. All Yesterdays is a philosophy that need only be applied to species for which soft-tissue data and behavioural aspects are inadequately known or entirely unknown. This includes some of the stranger Cainozoic mammals, reptiles and birds, and essentially everything that lived in the Mesozoic or before.

Composition, or aspect and attitude
What do extinct animals look like when not viewed in direct lateral view? Because the fossil skeletons of many species are laterally compressed, and because lateral views arguably show off more anatomy than other aspects, palaeoartists rarely show animals in anything other than side-on attitudes. 3D skeletal remains allow us to reconstruct animals in multiple aspects however, often with surprising and unfamiliar results. This has not percolated into palaeoart yet however, and if animals are shown in non-lateral aspects, their proportions are often 'generic'.  In my view, one of Greg Paul's crowing achievements was revealing the variation in dinosaur width, highlighting the extremely wide ribs of ankylosaurs and pachycepahlosaurs, and the narrowness of many theropods. All Yesterdays encourages us to remember that animals are three-dimensional beings, and can be accurately treated as such in art. (Below: the rarely seen anterior aspect of a famous pterosaur. But which one? Detail of a painting from my book.)

Appearance: the anti-shrink wrapping movement
Arguably the most important aspect of All Yesterdays is our reconsideration of extinct animal appearance. As a response to the inaccurate and often shapeless animals common to palaeoart in the early 20th century, most artists in the Age of Greg Paul employ the celebrated 'Rigorous Anatomical Approach' (RAA) to reconstructing fossil animals, and depict their animals with much of their detailed musculature and skeletal anatomy obvious under the skin (classic example of strict RAA in dinosaur art, Greg Paul's running Daspletosaurus, shown below. Image © Gregory S. Paul, from his website). These ultra-lean, toned animals are often devoid of any obvious extraneous tissues, including fats, loose skin or elaborate integuments. Strict RAA remains the most scientifically sound route to reconstructing an extinct animal, as it rigorously employs available data to render an extinct animal, and minimises speculation. It is also an effective way to demonstrate anatomical distinctions between extinct species and produces dynamic looking creatures, which are undeniably appealing to viewers. The rise in popularity of this technique at the time of the Dinosaur Renaissance is probably not a coincidence.

© Gregory S. Paul

However, most modern animals do not look like those produced under strict RAA. Fat, excessive skin, and integumentary structures hide much of their muscle profile and skeletal details, so they are not 'shrink wrapped' in the way that strict RAA suggests. The All Yesterdays philosophy embraces this fully, using RAA to provide the blueprint for an animal, but appreciating that not every anatomical feature will be discernible. Openings in the skull are not clearly seen on animal heads, large teeth are sheathed behind lips, limbs can be hidden beneath fur and feathers, details of muscles are obscured by wrinkly or thickened skin and so forth. Palaeoart produced with this in mind is more consistent with the appearance of modern animals, and make their subjects seem more plausible as living species. We must also not be too concerned about depicting extinct animals as looking ridiculous on occasion: there are numerous modern species which are frankly preposterous to behold, which surely must be true for some ancient species too. If our intended soft-tissue depictions can be functionally rationalised, and are consistent with fossil data and evolutionary hypotheses, then they are plausible inclusions for palaeoart. 

Behaviour
Anyone familiar with the history of palaeoart will recognise recurrent memes associated with specific animals. Ornitholestes always chases a bird. Archaeopteryx (which is always blue and green) perches on a branch with its wings outspread, its back always to the viewer. Tyrannosaurus is always roaring. Most prevalent of all is the depiction of prehistoric animals incessantly trying to murder each other. Memes often perpetuate because they reflect a certain trait specific to a certain animal (e.g. sleeping Mei, the use of the 'terrible claw' in Deinonychus), but they quickly became clichéd tropes when over used. All Yesterdays makes a case for showing animals undertaking other essential activities such as preening and bathing, socialising (without engaging in life-or-death intraspecific combat), playing, resting, sleeping, nesting and, well, all the other things that real animals do. Phylogenetic tracing of animal behaviour shows that there is no reason not to depict ancient animals undertaking these activities, but we rarely show them doing anything but fighting and eating. Moreover, All Yesterdays emphasises the behavioural plasticity of modern animals, noting that apparently strict carnivores or herbivores will supplement their diets with meat or plant matter on occasion, that animals can locomote in unexpected ways and are proficient at activities that we would not predict from their skeletons alone (this image being the chief All Yesterdays case study). As with the morphological aspects of reconstruction, this is not a call out for all unabashed craziness in palaeoart, but simply to say that we should be more broad minded about the way we depict the behaviour of extinct species, and that some initially outlandish ideas (like my carrion-eating Styracosaurus) are not as ridiculous as they first appear.

A new age in palaeoart?
Taken together, these points can be summarised in one two three sentences. Palaeoart must be both scientifically credible and realistic, but may be generally too conservative and clichéd to achieve these goals successfully. We are likely to be misjudging aspects of our reconstructions anyway, but we may be better off erring through the bold use of informed speculation about animal appearance and behaviour rather than through strict conservatism. Each produces results that cannot be refuted by current scientific data, but only the former produces art fully consistent with our understanding of real animals. 

It will be interesting to see how much of a shift All Yesterdays generates in attitudes to palaeoart. It's probably very clear by now that I'm a convert, but will others pick up on this, too? The rosy reviews of the All Yesterdays book suggest so, but what actual effect will this project have? I do not think the results will be as obvious as the popularisation of RAA in the 1970s and 80s, but I am optimistic that All Yesterdays marks a 'formalisation' of the anti-shrink wrapping movement, and need for depicting more complex compositions and behaviours. I look forward to seeing its results. Darren suggested at the All Yesterday's book launch that modern palaeoartists are currently working in the 'Age of Gregory S. Paul', with most of us following Paul's methods of strict RAA reconstruction to greater or lesser extent. Looking at the celebratory response to the All Yesterdays project across the internet, I wonder if the Age of Gregory S. Paul is about to end, and palaeoart will enter the 'Dynasty of All Yesterdays'?