Wesserpeton evansae: making 'albanerpetontid' a household name

Two Wesserpeton evansae get in each other's faces, because that's what albanerpetontids did. Note this is an updated version of the 2013 press release work referred to below. Prints of this image are available here.

You could be forgiven for thinking otherwise, but the Mesozoic wasn't just the remit of dinosaurs, pterosaurs, marine reptiles and token cool crocodiles. Many other interesting animals shared the world with these famous species, including some that most of us have never heard of. Tuesday of this week saw the (open access) publication of one such animal, the Wessex Formation albanerpetontid Wesserpeton evansae by Steve Sweetman (University of Portsmouth) and James Gardner (Royal Tyrrell Museum of Palaeontology) (2013). Many readers will be familiar with the Wessex Formation or the larger geological unit it is part of, the Wealden Supergroup, because of its frequent mentions as Britain's top dinosaur-bearing deposit. I'm sure many of us are not overly familiar with albanerpetontids, however. This isn't too surprising. To my knowledge, albanerpetontids have never featured prominently in any palaeoart or publications geared towards popular audiences and their existence is largely known only to specialists. The world's naivety to these animals was broken yesterday when Steve and James, with a little help from my painting above, finally told the world why they should add albanerpetontids to their list of cool fossil animals.

Alba-who?

Albanerpetontids are small-bodied amphibians that were fairly common components of terrestrial environments until relatively recently. The youngest members of their clan perished at the end of the Pliocene - about 2.5 million years ago - after an evolutionary run of 160 million years and attaining a wide geographic distribution across North America, Europe, Africa and central Asia. Their general lack of mention in popular press would have you believe otherwise, but they can actually be relatively common fossils. Remains of Wesserpeton are, after crocodiles, the most abundant microvertebrate in the Wessex Formation. Despite their relative abundance, their relationships to other lissamphibians have been debated because many of their fossils are exceptionally scrappy. Traditionally, they have been thought of as caudatans (salamanders) or at least very close relatives. Recent discoveries of complete and articulated albanerpetontid fossils (below) have suggested otherwise however, proposing that they are closely related to a clade containing frogs and salamanders, but not members of any extant amphibian group (McGowan 2002). 

LH 6020, holotype specimen of Celtedens ibericus, a complete albanerpetontid from Lower Cretaceous deposits of Las Hoyas, Spain. Note the 'halo' of scales around the fossil. Snout-vent length of this specimen is 59.5 mm. From McGowen (2002).

The anatomy of albanerpetontids is fairly conservative. They look more-or-less like small salamanders with short limbs and long bodies, but also possess mandibles which interdigitate anteriorly, fused frontals (bones of the skull roof) and relatively flexible necks because of a mammal-like articulation between the skull and neck. They also had bony scales under their skin, a condition which contrasts with the typically thin and delicate skin of most other amphibians. It seems that they spent most of their time burrowing through leaf litter in search of small arthropod prey, with their scaly skin possibly preventing dessication and likely reducing the typical amphibian need for wet or moist environments (but see comments below by the good David Marjanović). Fossils suggest that scales stretched across most of their bodies (we went the whole hog in our reconstruction and covered our Wesserpeton entirely) and onto their eyelids. We thought about these eyelids a fair bit for our painting. The few available depictions of albanerpetontids show animals with eyes perpetually covered with scales, leaving only very small, beady eyes to see with. Steve and I noted that these animals actually have very large orbits however, suggesting that their eyes were probably reasonably sized. It seemed counter-intuitive to possess large eyes and then cover them in soft-tissue, so our reconstruction assumes that the eyelids only partially covered the eyeballs.

Lower jaws of Wesserpeton evansae showing typical (A) and pathological (B) anatomies. From Sweetman and Gardner (2013). 

Small man syndrome

Initially, our plan for the press release painting was to show a single animal reclining in some leaf litter or something equally simple, but Steve suggested early on that we could work in an interesting component of Wesserpeton behaviour. Many Wesserpeton jaws show signs of trauma (above) after being broken during violent acts. The exact cause of this damage is still being looked into and will form the subject of a later paper, but a good preliminary explanation is that Wesserpeton was a vicious species which routinely fought among themselves. Modern salamanders, such as these giants, bite the heads of their opponents before wrestling with each other, twisting and somersaulting with one another to settle disputes over territory and mating access. It's not difficult to imagine such acts taking their toll on the jaws of Wesserpeton, and we thought it would be cool to show this in a press image. Preliminary attempts at rendering this struggled to show the general appearance of the animals however, as their bodies were twisted and their heads obscured by jaws. How could we show the aggressive nature of this animal without actually showing them fighting?

Do you speak salamander? Common body postures used to communicate between individuals of the red-backed salamander Plethodon cinereus. We took panel B as our primary inspiration for the Wesserpeton evansae PR image. Figure and caption from Jaeger (1984).

The solution came in the form of postural language borrowed from modern red-backed salamanders (Jaeger 1984, and above).Most animals will attempt to intimidate their rivals with ritualised postures which enhance their perceived size and strength before coming to physical blows, and there's no reason to think that little albanerpetontids were any different. We decided to use the postures of modern salamanders in our image, setting one of our animals as a dominant pose with a raised trunk and tail, and the other crouched and submissive. In doing so, we implicated the violent nature of this species (enhanced by the larger animal getting in the face of the smaller, just like most douches with attitude problems) but maintained the ability to show their anatomy. Entirely coincidentally, Darren Naish and John Conway recently spoke about incorporating animal postures into palaeoart in Tetrapodcats (sic) episode 6, which makes for interesting listening if you're thinking about making your restored extinct animals communicate more fluidly.

Finally, a quick word on the body size of Wesserpeton. We've mentioned it was small, but how small? The answer is tiny. As in, 35 mm snout-vent length tiny. This thing really puts the 'micro' in 'micropalaeontology'. We prepared another set of press images to show what this means in real life (available in different colours to suit whatever occasion you're at where you want to discuss the size of Wesserpeton):

The United Colours of Wesserpeton, which is dwarfed by the palm of your hand no matter what colour you are.  For some reason, this image makes me want to imagine a world without lawyers.

I'm no expert on this sort of thing, but I'll wager that Wesserpeton is one of the smallest, if not the smallest tetrapod species known from the Wessex Formation, and probably one of the smallest tetrapods in the fossil record. It's fossils were only recovered through bulk sampling tonnes of plant debris bed from the Wessex Formation, horizons rich in plant and vertebrate remains deposited after sheetflood events, and would be almost impossible to find via surface prospecting. Those of you with excellent memories may recall that Steve's ongoing analyses of these beds have revolutionised our understanding of the Wessex palaeobiota, of which Wesserpeton is just one discovery among many. 

And that will have to do for now. Next week: back to the world of pterosaurs with pterosaur mummies, as promised last week before Wesserpeton face-wrestled its way into centre stage. My plan from here on is to have some sort of run-up to the publication of my book, Pterosaurs: Natural History, Evolution, Anatomy on June 23rd, so be sure to stick around if wing membranes are your thing.

References

• Jaeger, R. G. 1984. Agonistic behavior of the red-backed salamander. Copeia, 309-314.
• McGowan, G. J. 2002. Albanerpetontid amphibians from the Lower Cretaceous of Spain and Italy: a description and reconsideration of their systematics. Zoological Journal of the Linnean Society, 135(1), 1-32.
• Sweetman, S. C., and Gardner, J. D. 2013. A new albanerpetontid amphibian from the Early Cretaceous (Barremian) Wessex Formation of the Isle of Wight, southern England. Acta Palaeontologica Polonica, 58, 295–324.

Another Pterosaurs preview, and the soft bits of Tupandactylus

Tupandactylus navigans reclining by sunset, pycnofibres a-glowing.

Holy Toledo, the publication date of Pterosaurs: Natural History, Evolution, Anatomy is now only weeks away. In exactly one month, preordered copies of the book will be sent out and actual, physical copies of it will be in homes around the world. Forgive me if this sounds indulgent: it's not meant to. It's simply a little mind boggling to think of people paying hard-earned money for the book that, with help from Princeton University Press, I spent over two years writing and illustrating. (Don't forget to add the celebratory Pterosaurs party at the Natural History Museum, London, on September 10th, 2013, to your diary.)

To celebrate this navel-gazing milestone, here's another preview image from the book. It shows the Brazilian tapejarid Tupandactylus navigans at sunset, it's fur-like pycnofibres glowing in the diminishing light. This painting is one of the large paintings that accompanies the start of each chapter and, specifically, it opens Chapter 5: "Soft bits". Each of these large paintings was designed to draw focus to the topic of its chapter. Deciding on the basic composition was easy enough for many chapters, but those focussing on soft-tissue anatomy and osteology proved to be a little bit of a head scratcher. How do you draw specific attention to tissues comprising pterosaur bodies rather than the pterosaur itself? The answer for "Soft bits" at leastseemed to lie in back lighting a pterosaur body so that most of the animal was obscured, save for a halo of illuminated fuzz. Tupandactylus navigans was chosen because it's enormous soft-tissue headcrest (below) contributed to the already unusual outline of a pterosaur body to make a more startling image. "Soft bits" takes on a variety of other soft tissues as well - brains, lungs, guts, skin, wing membranes and so forth - but these seemed harder to bring out without cutting a pterosaur open.

Tupandactylus navigans holotype skull SMNK PAL 2344, showing the crazy headgear sported by some tapejarid species.  Remember that this crest is not the largest worn by a tapejarid pterosaur. From Witton (2013).

In other news, today also saw Christopher DiPiazza post an online interview he conducted with me at Jersey Boys Hunt Dinosaurs. The post contains some exclusive artwork, comments on how to get into palaeontology and a frank reply to the question of "should I undertake postgraduate studies in palaeontology?" I'm not the only chap telling people to be cautious about choosing palaeontology as a career at the moment, and seriously recommend that prospective students thinking about joining the palaeo ladder give that choice some serious thought before taking the plunge. Read why here (question 7).

That's all for this week. Next week: pterosaur mummies!

P.S. 'Tupandactylus' navigans? Who he?

Finally, a quick note on the nomenclature used in this post. Readers familiar with tapejarid taxonomy may notice that I'm treating navigans as part of the genus Tupandactylus, whereas it has typically been referred to Tapejara or "Tapejara" by other workers. The nomenclatural history of navigans is a little complicated. It was initially placed in the genus Tapejara (Frey et al. 2003) along with two other species, T. wellnhoferi and imperator. Two teams of authors independently revised the taxonomy of this genus in 2007, with Kellner and Campos (2007) moving imperator to a novel genus, Tupandactylus and Unwin and Martill (2007) creating another new genus, Ingridia, for navigans and imperator, with the latter as the type species. The work of Kellner and Campos was published just before Unwin and Martill and, because they both used imperator as the type taxon of their respective genera, Ingridia must be considered synonymous with Tupandactylus. Kellner and Campos (2007) hinted that navigans was also probably a member of Tupandactylus, but Darren Naish suggested that it may still warrant generic distinction from imperator in a 2008 Tetrapod Zoology article. navigans has been in taxonomic limbo since then, but recent phylogenetic work (e.g. Pinheiro et al. 2011 and my own studies, presented last year at SVPCA 2012 and hopefully being turned into a fully fledged paper when I get the time) has found support for a navigans + imperator clade which bears out earlier suggestions that these species are congeneric. These discussions about generic labels are fairly arbitrary and someone may eventually decide to generically split navigans from tupandactylus but, until then, it seems reasonable to house navigans within the Tupandactylus stable. 

References

• Frey, E., Martill, D. M., and Buchy, C. C. 2003.  A new species of tapejarid pterosaur with soft tissue head crest. In: Buffetaut, E. and Mazin, J. M. (eds.) Evolution and Palaeobiology of Pterosaurs, Geological Society Special Publication, 217, 65-72.
• Kellner, A. W. A. and Campos, D. A. 2007. Short note on the ingroup relationships of the Tapejaridae (Pterosauria, Pterodactyloidea). Boletim do Museu Nacional, Nova Séroe, Rio de Janeiro - Brasil. Geologia, 75, 1-14.
• Unwin, D. M. and Martill, D. M. 2007. Pterosaurs from the Crato Formation. In: Martill, D. M., Bechly, G. and Loveridge, R. F. (eds) Window into an ancient world: the Crato fossil beds of Brazil, Cambridge University Press, Cambridge, 624 pp.
• Pinheiro, F. L., Fortier, D. C., Schultz, C. L., De Andrade, J. A. F., and Bantim, R. A. 2011. New information on the pterosaur Tupandactylus imperator, with comments on the relationships of Tapejaridae. Acta Palaeontologica Polonica, 56(3), 567-580.
• Witton, M. P. 2013. Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. [In press]

Another new Plateosaurus

A modern reconstruction of Plateosaurus engelhardti as a relatively bird-like, bipedal animal rather than a tubby, sprawling  quadruped

It turns out that you can teach an old dinosaur some new tricks. Remains of Plateosaurus engelhardti have been known since at least 1837 but, thanks to a flurry of recent research into its anatomy, posture and gait, we've learnt many surprising facts about this animal in the last few years. The restoration offered here is an attempt to portray this animal in its most recent, modern light and, in many details, it contrasts markedly with reconstructions even only a few years ago.

'New looks' for Plateosaurus are not rare, however. Our perceptions of this dinosaur have been rather changeable since its discovery, and few dinosaurs have been through as many iterations of posture, gait and general appearance. Since its discovery in the early 1800s, we've seen Plateosaurus reconstructed in numerous ways. Perhaps the oldest known reconstruction, by O. Jaekel, dates to 1913-1914 and depicts a very different animal to the one we know today. This frequently proportionally inaccurate image shows a very odd-looking bipedal dinosaur which almost has a therizinosaur-like appearance, but perhaps is better compared to Igor from an early Universal horror film. Later depictions of Plateosaurus were of a sprawling quadruped with a barrel-like chest; an erect biped with a dragging tail; a lithe quadruped capable of galloping; a very sauropod-like animal with straight, columnar limbs and a long, low biped with a heavy balancing tail. These interpretations, and others, have been compiled and thoroughly discussed by Heinrich Mallison, a man who clearly loves Plateosaurus as much as a man can love a fossil species (below, from Mallison 2010a).

The many faces of Plateosaurus, compiled by Mallison (2010a). Check out Heinrich's open-access publication for sources of the reconstructions.

This wealth of conflicting interpretations is odd when we consider that, among non-sauropodan sauropodomorphs and perhaps dinosaurs generally, the osteology Plateosaurus is very well known. Numerous articulated skeletons, some essentially preserved standing in miring muds, inform us about details of limb carriage and girdle configuration (see image, below), but it seems that much of this data was overlooked by scientists and artists restoring Plateosaurus for decades. Once the fine anatomy and taphonomic data of these animals was assessed in detail however, it became very apparent that most of our interpretations of Plateosaurus were simply plain wrong. The palms of the Plateosaurus hand were forever facing medially, like those of theropod dinosaurs and unlike many of the reconstructions shown above (Bonnan and Senter 2007). It's forelimbs had an extremely limited range of motion, which more-or-less only permitted movement beneath the body (Mallison 2010a). When the entire skeleton was reconstructed without disarticulated bones, the narrowness of the chest and shoulders became apparent, contrasting with the enormous bellies of many mounted skeletons and sculptures. The rigidity of its avian-like torso skeleton and disproportionate limbs were also brought to light (Mallison 2010a, b, and references therein).

SMNS F33, one of the most famous articulated fossils of Plateosaurus engelhardti. Note the narrowness of the chest and shoulders. Photograph by Heinrich Mallison, uploaded to Wikipedia Commons by FunkMonk.

In essence, it seemed that virtually all reconstructions of Plateosaurus prior to 2010 were committing some sort of major anatomical sin, requiring disarticulation of bones or over-flexed joints in order to achieve their portrayals. Perhaps most obviously, it seems that quadrupedal locomotion, be it a slight sprawl or a horse-like gallop, was not possible for Plateosaurus (and perhaps most other non-sauropodan sauropodomorphs). Rather, the reach, attitude and length of their forelimbs indicate that they were habitual bipeds, and their centre of gravity dictated that they held their backs subhorizontally. Gone too were the round, barrel-like chests: despite being a  6-10 m long herbivore, Plateosaurus was a surprisingly gracile animal that was perhaps even capable of moving relatively quickly on its powerful back legs. The contrast of this new appearance with some of the older, sprawling reconstructions of this animal is rather pronounced.

(Interesting bonus knowledge to stem from recent Plateosaurus studies include new insights into the suppleness of the Plateosaurus neck, which permitted 360° to their owners. Presumably, this reflects a need to keep an eye out for predatory archosaurs which, in the Late Triassic landscapes occupied by Plateosaurus, would not be predatory theropods but large rausuchians like Teratosaurus. [How come there's not many illustrations of predatory interactions between these species? Would probably make for a cool image]. Perhaps even more interestingly, the pedal claws of Plateosaurus show evidence of being used in scratch digging, which may reveal how these animals dug their nests. Neat stuff indeed.)

Junk about the trunk
But it's not just Plateosaurus specific topics that have fuelled the latest changes in its appearance. It's very likely that the tail of Plateosaurus was more massive and heavy than portrayed in most reconstructions because, along with many other dinosaur species, it bore substantial hindlimb musculature along the base of its tail. We can see how large the hindlimb retractor muscles of dinosaurs should have been by looking at homologous anatomies in lizard and crocodile tails (warning: that link is not for the squeamish), and the take-home message is that most dinosaurs had extremely beefy proximal tail regions (e.g. Persons and Currie 2011). As Heinrich covers here, we can see osteological evidence for expansive, tail-anchored hindlimb muscles in Plateosaurus as clearly as we can in nearly all other Mesozoic dinosaur species. The Plateosaurus anterior tail regions was effectively a big tube of meat bulging from a thread of vertebrae, which were themselves enlarged for the attachment of powerful muscles. In life, the hindlimb protractors would probably run continuously from the tail to the hindlimb, with no obvious 'join' between these two body sections. These bulging tails are a world away from the slender and compressed tails seen in many pieces of palaeoart. Furthermore, all this proximal tail muscle probably reduced the flexibility of the tail somewhat (Persons and Currie 2011) suggesting that another palaoart trope - dinosaurs with laterally wavy, sinuous tails - were probably impossible. The extensive muscles at the base of the tail can only be compressed so far by a wiggling tail. Thus, although the tail bones of Plateosaurus seem to offer a fair amount of lateral motion (below; also see Mallison 2010a), there probably wasn't anything close to that level of flexibility available in life. In the picture above, the tail is deliberately a little lifeless, acting purely as a site for hindlimb retractor muscle anchorage and a counterbalance to the heavy, gut-filled torso.

Digital model of Plateosaurus caudal vertebrae in dorsal view, showing 10° flexion between each joint. From Mallison (2010a).

Feathers? Really...?
So, new posture, new build, new tail, and now new integument? We're very used to seeing Plateosaurus and with virtually all other sauropodomorphs with scales, but this may not have been the case  (hat tip here to recent blog posts on this topic by the Internet's favourite sauropod workers, Matt Wedel and Mike Taylor). Sauropodomorphs may not have provided any fuzzy revelatory fossils over the last few years but, because lots of other dinosaur groups have, the landscape of dinosaurian integument has shifted considerably around their feet. Nowadays, it's a commonly known possibility that dinosaurs, and perhaps all ornithodirans, were ancestrally fuzzy. Although we have a few skin impressions from sauropods (Czerkas 1992; Coria and Czerkas 2007), they remain pretty rare across the group as a whole and, to my knowledge, we don't have any from non-sauropodan sauropodomorphs. Our few scraps of sauropod skin suggest they bore typically archosaurian, non-overlapping scales but, even if that's standard for the group (and, given what we know about modern animal integument, it may well not be), it isn't clear when this trait became a typical part of the sauropod bauplan. Thus, it is not inconceivable that some early sauropodomorphs were feathered and, until we have some direct evidence either way, it's probably just as parsimonious (and, arguably, speculative) to draw them with fuzzy hides as it is with scales. In the case of the picture here, I figured throwing an ostrich-like distribution of feathers over a modern Plateosaurus reconstruction added a suitably symbolic purpose to show how far interpretations of this animal have come since its first discovery.

And that may be enough novelties for one day: time to crack on with other things. Here's hoping that this reconstruction of Plateosaurus will stand the test of time, though I'm sure I'm merely tempting fate with those words. Next week, hopefully, we'll see something to do with animals bearing membranes, because this thing isn't very far from being released on the world any more.

References

• Bonnan, M. F., and P. Senter. 2007. Were the basal sauropodomorph dinosaurs Plateosaurus and Massospondylus habitual quadrupeds? In: Barrett, P.M. and Batten, D.J. (eds.), Evolution and palaeobiology of early sauropodomorph dinosaurs. Special Papers in Palaeontology, 77, 139-155.
• Coria, R. A. and Chiappe, L. M. 2007. Embryonic skin from Late Cretaceous sauropods (Dinosauria) of Auca Mahuevo, Patagonia, Argentina.  Journal of Paleontology, 81, 1528-1532.
• Czerkas, S.  A. 1992. Discovery of dermal spines reveals a new look for sauropod dinosaurs. Geology 20, 1068-1070.
• Mallison, H. 2010a. The digital Plateosaurus II: an assessment of the range of motion of the limbs and vertebral column and of previous reconstructions using a digital skeletal mount. Acta Palaeontologica Polonica, 55,433-458.
• Mallison, H. 2010b. The digital Plateosaurus I: body mass, mass distribution and posture assessed using CAD and CAE on a digitally mounted complete skeleton. Palaeontologia Electronica 13.2.8A
• Persons IV, W. Scott, and Philip J. Currie. 2011. The tail of Tyrannosaurus: reassessing the size and locomotive importance of the M. caudofemoralis in non‐avian theropods. The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology, 294.1, 119-131.

Ray Harryhausen, 1920-2013

A prehistoric animal being lassoed by cowboys alongside ruined temples in a barren landscape? Sounds like a Harryhausen movie. Oh wait: it kinda was.

There aren't many people in the public eye who I admire, but the pioneering stop-motion artist Ray Harryhausen was one of them. Harryhausen didn't kick start my interest in palaeontology in the first place, but his animated dinosaur sequences were certainly one of the highlights of my dinosaur obsessed childhood. I still watch his films today, both dinosaur-themed and otherwise. In fact, I watched two of them just this weekend. Turns out this was strangely timely: as we all know by now, it was announced that Harryhausen passed away yesterday (7/5/2013) at the age of 92.

I'm not alone among palaeontologists in revering Harryhausen's work. Few other special effects artists in the 20th century could make dinosaurs and other prehistoric animals come alive quite like Harryhausen, and it was perhaps only the sophisticated animatronics and CG effects in 1993's Jurassic Park which clearly superseded his work. Indeed, to appreciate Harryhausen's work fully you had to live in pre-JP times, where movie dinosaurs were generally pretty poorly executed. Before Jurassic Park, most movies dinosaurs were modern lizards and crocs dressed up with horns and sails, shonky animatronics and puppets, or else men in oversize reptile costumes. None of these approaches were terribly convincing. Harryhausen's stop motion models, by contrast, looked and acted like the real thing, battling with people and other creatures as if they were present on set. With very few exceptions, there was simply nothing closer to seeing actual dinosaurs than Harryhausen's animations until 1993. It's hard to chose a favourite scene from his dinosaur works, but in terms of overall execution, I think his Allosaurus attack in the 1966 film One Million Years BC may be among his best (below, borrowed from Apollomovieguy). As with all Harryhausen's creations, the animal in this scene is full of character thanks to subtle movements of its head and body. At times the allosaur comes across as a genuine actor in the production, and one who thinks that the movie is below him. Some of his lazier snaps and turns suggest it's just walking the part rather than playing the ferocious animal it could be. You can almost see it thinking 'well, at least I'll be able to pay the rent this month' as it snaps, idly, at the cavefolk. It's only when it's allowed to run across the set and savage people that it really seems to surrender to the role. I don't mean these a criticisms: Harryhausen's careful animation that gives the animal these little flourishes of character that make his films a joy to watch. There are numerous other examples we could cite of this sort of thing: Talos staggering a little when climbing from his plinth, and is preferentially uses his right hand, in Clash of the Titans (1981); Gwangi tugging at the lasso ropes around his neck with his little arms in Valley of Gwangi (1969) and so forth.



Although we may automatically think of dinosaurs when Harryhausen is mentioned, he actually only made three movies which contain 'proper' Mesozoic dinosaurs: The Best from 20,000 Fathoms (1953), One Million Years BC (1966) and Valley of Gwangi. Indeed, we're only including Beast on a technicality. The plot suggests that the 'Rhedosaurus' creature is a dinosaur, but it looks more like a giant lizard than a theropod. I get the impression that Harryhausen would have liked to have worked with dinosaurs more,  apparently trying to drum up interest for further other dinosaur movies or crowbar them into other productions (Sinbad was meant to meet a ceratopsian in a fourth film in that series). He certainly seems to frequently mention dinosaurs in a favourable light in interviews, and some of his first stop-motion models - made when he was a young teenager, were of famous dinosaur species. He did, of course, animate a short sequence of dinosaurs for the 1956 documentary Animal World, which was his only attempt to show dinosaurs in a natural time and setting*. A few other Harryhausen movies - Mysterious Island (1961) and Clash of the Titans - featured avian dinosaurs, but, in all, he actually made relatively few productions featuring the 'classic' dinosaurs. Much of his filmography is comprised of other fantasy or science fiction fare, so his dinosaur movies are, in fact, a minority of his work. Clearly, his dinosaur scenes were well executed enough to remain prominent in our memories even if, when put together, they only comprise a few minutes of footage.

*Unlike his other dinosaur scenes, the models for Animal World were made from injection foam, a cheaper but inferior material to his preferred medium of layered latex, and the sequence features closeups of awkward looking robotic miniatures. For this and other reasons, Harryhausen was never really happy with his work on Animal World and, although the stop-motion still stands up, it's easy to see why he did not consider it as highly as his other projects.

I find it interesting that palaeontologists are so fond of Harryhausen's work considering his sometimes flippant regard for scientific accuracy. Palaeontologists can be real sticklers for such issues, and Harryhausen had a pretty loose concept of what dinosaurs were like (which may explain his 'dinosaur' in The Beast from 20,000 Fathoms). He frequently over-scaled his animals to make them more formidable, or hybridised different species for dramatic effect. The theropod Gwangi is well known as being a hybrid of Tyrannosaurus and Allosaurus, and he added bat-like wings and owl talons to pterosaurs (see his Pteranodon model, below, borrowed from my Pterosaur.Net post on this very topic). Harryhausen played loose with myths as well. His harpies, featured in the 1963 film Jason and the Argonauts, have bat-like wings instead of the feathered wings described in folklore, and his Medusa, from Clash of the Titans, sports a rattlesnake abdomen not mentioned in any literature on this creature.

Despite his loose interpretation of source material, Harryhausen rarely invented completely novel anatomies for his creatures, even when working with entirely novel concepts. His anatomies always had a source in the real world, and it was his combination of animal components which resulted in different models. It's for this reason that Harryhausen's creations share the same appearance, as some of his anatomical 'building blocks' have a signature style. The Kraken from Clash of the Titan, for instance, has the body resembling his JatA Talos model (down to the navel and nipples, which are curious features on a titan), the head is rather like Ymir from 20 Million Miles to Earth (1957) and the arms are clearly borrowed from the octopus in It Came from Beneath the Sea (1955). These characters all represent very different types of beast - mythological titans, aliens, and giant animals - but they all share Harryhausen's artistic DNA.

For all of his recycling of designs and body parts though, Harryhausen never tweaked his creations so far from their source material that they were unrecognisable. They were enhanced, sure, but never distorted. Palaeontologists watching his dinosaur flicks could feel satisfied that his animals more-or-less looked matched concepts of these animals fashionable in the 50s and 60s. Besides, his animations are too unapologetically focussed on entertainment to warrant harsh scrutiny. You can nitpick Gwangi's anatomy all you like, but you'll be missing the coolest dinosaur round up scene ever committed to film if you do. The plots of Harryhausen's films, which he was often a key producer of, are all about pulpy entertainment and nothing more. They appeal to the little boys and girls in all of us, and aren't asking for detailed critiques of their stories, scientific plausibility or even scripts and acting. Can anyone in touch with their inner child honestly say that they're not interested in watching cowboys round up a tyrannosaur, or a dinosaur destroying a lighthouse? Harryhausen's ideas are simply so charismatic that they're practically immune to scientific criticism.

Harryhausen's legacy is not just one a charismatic storyteller and skilled special effects artist, however. It's well known that Harryhausen was one of the signature stop-motion artists of the 20th century, but the fact that he saved stop-motion as an artform in the 1940s and 50s isn't common knowledge. Early stop-motion techniques pioneered in films like The Lost World (1925) and King Kong (1933) by Harryhausen's mentor, Willis O'Brian, were wonderfully executed but extremely time consuming and expensive, even by standards of stop-motion photography. O'Brian's techniques employed setting several painted landscapes on glass between his animated models and the camera, thus creating the impression that the creatures were set in middle distance. The process of painting these landscapes and compositing these shots, in addition to animating the models and other technical work, was so intensive that most studios were unwilling to invest the necessary time and money into stop motion work. Harryhausen's childhood interest in stop-motion animation, fired by multiple viewings of King Kong and some very supportive parents, allowed him to develop matting techniques which negated the need for painted glass landscapes, in which components of the filmed footage could be re-photographed on top of the animated scene. This cut the time and material costs of stop-motion processes drammatically, and he pioneered this technique, which he would eventually call 'dynamation', before he landed his first solo job on a motion picture. Dynamation was first put to the test on the low budget Beast from 20,000 Fathoms, and Harryhausen delivered his work for a fraction of the cost of traditional stop motion. Without this revolution in stop-motion, the artform as we know it may never have happened. That may not seem like a big deal to us now in a time of CG effects, but bear in mind that Jurassic Park was greenlit with a sophisticated form of stop-motion in mind.



In his career, Harryhausen more-or-less single handedly lent his dynamation craft, and other special effects wizardry, to something like 16 motion pictures (see compilation, above, by Vidar Solaas). If the testimonies of the numerous fans and film makers are accurate, the movie world would be a very different place without them. There's so much more we could say about Harryhausen: his abilities as an artist and sculptor, the vast numbers of films he designed that were never made (War of the Worlds with Harryhausen tripods!), the many, many stories he has shared about making his films, but there's simply not enough time to relate even a fraction of them here. If you want to know this sort of stuff, though, I thoroughly recommend you track down either his richly illustrated autobiography and collected artworks, or the recent critically acclaimed documentary Ray Harryhausen: Special Effects Titan.

On that note, it's time to get on with other things. Thanks then for everything, Ray, you will be missed.

The Walking with Dinosaurs 3D trailer lands

Good news, everyone: the PR machine behind Walking with Dinosaurs 3D has provided our first detailed look at the film with its first trailer. I'm rather excited about this because it's my first real look at the film since contributing character orthographics for several (six, specifically) of its animals in early 2011, a process that involved working with a fleet of palaeontological and palaeoartist supernauts: David Krentz, Scott Hartman, Tom Holtz, Scott Sampson, Luis Chiappe, Victoria Arbour and others. Since that time, the guys at Animal Logic have been crunching the ones and zeroes required to breath life into the characters and their world, with the results posted above. There's always an air of uncertainty surrounding palaeontological film projects but, happily, this first trailer suggests WwD3D is taking good shape, and the Facebook and Twitter buzz from the contributors mentioned above suggests they're pretty happy with what they've seen, as am I. Be sure to set the trailer stream to its highest, 1080p HD quality to take in all the details.

Highlights of this first teaser, aside from some excellent zbrush creations from David Krentz and the stunning visuals and animation, include feathered dromaeosaurs, an iridescent gorgosaur and suitably snowy slopes in latest Cretaceous Alaska. I like the efforts to characterise individual animals without resorting to crazy variations in colour and form, such as the hole in the frill of the hero Pachyrhinosaurus. Presumably, this reflects a pathology that never entirely healed, and it's a cool idea. So cool, in fact, that I'm wondering why it's not more common in palaeoart. We can also see that the colour schemes of the animals are striking without being outlandishly garish, and the characters seem capable of emoting without using overblown actions akin to those of other CG dinosaurs, who seem to have gone to an acting school for 1920s silent movies. Moreover - stop press - plenty of shots even show animals not roaring and with their mouths closed. Gosh.

So, there's plenty to be happy about then, and my feeling from reading the comments of others is that I'm not alone in this view. Perhaps the only common niggle being raised is that the tyrannosaur is scaly rather than feathered, which jars with recent discoveries showing some big tyrannosauroids were covered in protofeathers (Xu et al. 2012). Obviously, it would have been extremely cool to render this in WwD3D, but the feathered tyrant Yutyrannus arrived just a little too late in the day for the design team to work feathers onto the Gorgosaurus. As hinted above, the creature designs for the WwD3D animals were being set in late 2010 and early 2011, well over a year before Yutyrannus was unleashed on the world. And we shouldn't be too upset: there is still controversial evidence that another tyrannosaur, Tyrannosaurus, was scaly, and the WwD3D Gorgosaurus still looks awesome.

But enough about dinosaurs: what about the real stars of the show, the pterosaurs? It may not be surprising that I had a hand in design and consultancy for the movie's azhdarchids, and I'm happy to say that I like what I can see thus far. Flap-gliding and full-on flapping flight are both on display, and just short of the one minute mark, we see a gang of azhdarchids surround and aggressively 'terrestrially stalk' the hero animals. If you're sad enough to have freeze-framed high-definition shots of the azhdarchids (which, er... I didn't, but a friend of mine did) you'll notice thick coats of soft pycnofibres, and that the internal anatomy of azhdarchid jaws have, for the first time, been accurately rendered on film. Indeed, the contours of the head are the best I've ever seen in a CG azhdarchid. Note the concave lateral skull facia, the tapering shape of the tomial margins... wonderful stuff. I'm very much looking forward to seeing them on screen later in the year.

So, this is all shaping up very well then, making the December 20th release date something to look forward too. It almost seems that fans of Mesozoic reptiles on film are being spoilt at present, with WwD3D following 2011's terrific (and underrated) Dinosaur Revolution/Dinotasia and the excellent BBC series Planet Dinosaur. Indeed, the BBC's dinosaur coverage is on something of a roll at present. Presumably as part of the PR campaign for the new film, its revamped Walking with Dinosaurs website has been providing good coverage of recent dinosaur topics in a way that is extremely accessible but not dumbed down (for a good example of this, see their coverage of the Torosaurus/Triceratops debate). Several well-known contributors to the palaeo blogosphere have been contributing new content to the site too, so it's definitely worth checking out if you haven't already.

And that will have to do for now: lots of work to catch up with. In the meantime, I leave you, and particularly if you're a 20th Century Fox or BBC executive, with a pitch for a WwD3D spin-off, starring the animals I'm sure will steal the entire movie. It's Box Office Gold, I tell you.

Reference

• Xu, Xing, Kebai Wang, Ke Zhang, Qingyu Ma, Lida Xing, Corwin Sullivan, Dongyu Hu, Shuqing Cheng, and Shuo Wang. 2012. A gigantic feathered dinosaur from the Lower Cretaceous of China. Nature, 484, 92-95.