Review: "Beasts of the Mesozoic" Tsaagan by Creative Beast Studios

Quick Facts
2018 Beasts of the Mesozoic Raptor Series Tsaagan mangas action figure
Size: 20cm long
Scale: 1:6
Sculpted by: David Silva
Produced by: Creative Beast Studios

Back in April 2016, toy industry veteran David Silva launched a Kickstarter campaign to produce scientifically accurate "raptor" (eudromaeosaur) figures. Unlike the vast majority of static PVC dinosaur figures on the market, these would be super articulated, with up to 24 joints allowing significant posability. Now, over two years later, the project has become a reality, and my selection of a Wave 2 Tsaagan mangas figure has finally arrived. So, how does it stack up to the high expectations and lofty claims that these are the most scientifically correct dinosaur action figures on the market?

Review: Dino-Riders Struthiomimus by TycOMG IT HAS FEATHERS

Quick Facts
1988 Dino-Riders Struthiomimus action figure WITH FEATHERS. IN 1988.
Size: 20cm of feathered glory.
Scale: Scales on the feet, feathers up top. Also, 1:12.
Sculpted by: The wokest of all 1980s dinosaur toy sculptors.
Produced by: Tyco (obviously with a lot of help from Bob Bakker).

No need to adjust your TV sets folks, this is a mass-produced dinosaur toy made in 1988 that is covered in feathers. Not like lame, Primal Carnage, Jurassic Park 3, cool-guy dragon with a mohawk. Natural looking feathers.

This is why Dino-Riders was the best thing about the '80s (sorry, He-Man). Dino-Riders gave us aliens from the future riding armored mind-controlled dinosaurs blasting a thousand lasers at other armored dinosaurs who were not mind controlled but who were just in it because they cared about justice, and the toy versions of these things looked more naturalistic and scientifically accurate (for the time) than anything in Jurassic World.

I'm going to use this particular review to drop some history.

You're Doing It Wrong: Microraptor Tails and Mini-Wings

Type specimen of Zhenyuanlong, doing its best Archaeopteryx impression.

Just a short PSA today, and once again, it's about a paleoart meme that has outstayed its welcome.

Microraptor was the first time we got a good look at the feather pattern of dromaeosaurids. This is a big problem for two reasons. One, microraptors were small. That means that artists who were looking at them to extrapolate for bigger, more famous "raptors" could easily and somewhat justifiably write off their huge wings as a product of their size. Sure, we thought, microraptors had big wings, but they're tiny animals. Surely the bigger, more terrestrial dromaeosaurids didn't need such big wings. They probably still had wings, but they'd be smaller. Why would Velociraptor need such proportionately huge wings if it couldn't fly or glide?

Meme number two: that tail. I admit to being one of the first to go overboard when I fell, head over heels, for the "puff tailed dromeosaur" fossil (now the holotype of Cryptovolans, a synonym or close relative of the Microraptor) back around 2000. This was the first evidence we had of the tail feather style in dromeosaurids (or evidence that they even had remixes and rectrices at all. Remember When Dinosaurs Ruled America? That was plausible at the time it was being made). Naturally, having Microraptor plus Caudipteryx showed that the ancestral condition of pennaraptorans was a fan of feathers at the tip of the tail, not a fuzzy Sinosauropteryx like tail or a fully-vaned Archaeopteryx like tail. So artists ever since have been drawing dromeosaurids and troodontids and oviraptorosaurs with microraptor tails.

But that turned out to be wrong! It's an accident of history. We're now learning that Microraptor and Caudipteryx are weirdos.

The First Feathered Dinosaurs (In Art)

The first illustration of a hypothetical "pro-avis" by Pycraft, 1906

Feathered non-avialan dinosaurs seem commonplace now, and it's hard to believe that there was a time, in living memory for some of us, when they were purely speculative. It makes sense, of course: once scientists realized (then realized again) that dinosaurs were the closest fossil relatives of birds, it was only natural to suggest that feathers had appeared in dinosaurs before they showed up in birds. To have a new type of animal, like Archaeopteryx, appear more or less fully formed with flight and feathers occurring simultaneously makes little sense in terms of evolution. Obviously, even if Archaeopteryx was considered the "first bird" or could fly in some rudimentary manner, it couldn't have been the first animal with feathers or feather-like integument. Feather-like structures had to have been present in the archaeopteryx's closest relatives. But what were those? For most of the 20th century, the answer would not have been dinosaurs.

Nopcsa's 1907 feathered, Compsognathus-like
"pro-avis" model at the Grant Museum, London.

Speculative bird ancestors with feathers or feather-like scales had been hypothesized and illustrated long before the first true feathered dinosaurs appeared in paleoart. These hypothetical "pro-aves", as they were generally termed, were first imagined by William Pycraft in 1906. Pro-aves were usually considered arboreal, gliding animals with elongated, feather-like scales, and Pycraft's illustration provided the template followed by many later illustrators, like Heilmann's 1916 version, and Burian's gorgeous but even more lizard-like 1960 rendition. Not all pro-avis illustrations followed this template, though. In 1907, Baron Nopcsa invented his own pro-avis, illustrating his hypothesis of a running, "ground-up" origin of bird flight. In addition to his illustration, Nopcsa created a wax model. Nopcsa thought birds might have evolved from quadrupedal reptiles which became more and more bipedal as their running speed increased, passing through a phase similar to Compsognathus, though not necessarily dinosaurian.

Beebe's "tetrapteryx", 1915.

There were also creatures like the "tetrapteryx", a speculative stage in the evolution of avian flight proposed by William Beebe in 1915. Beebe illustrated an Archaeopteryx-like creature with smaller wings as well as hind wings on splayed legs, reminiscent of early (but incorrect) illustrations of Microraptor. Other than the posture, the general anatomy of this hypothetical creature ended up being prescient by accident. His rationale for this hypothesis was the observation that some bird embryos develop and then lose feather quills on their legs. Heilmann, however, rejected this hypothesis after failing to find evidence among other bird embryos. New evidence to support this hypothesis was discovered in species like Microraptor, Anchiornis, and Sapeornis, which had various degrees of airfoils present on their hind legs, a line of evidence independent of the one supposed by Beebe. Though dinosaur-like in appearance, both Beebe and Heilmann considered the ancestors of birds to be "thecodont" grade archosaurs, not dinosaurs, so these hypothetical feathered thecodonts predated the first feathered dinosaurs in art.

Speculative reconstruction of a Triassic "carnavian" dinosaur
leaving feather traces in its trackway, from Ellenberger 1974.

The thecodont origin of birds did not truly give way to a dinosaur origin until the 1970s, after John Ostrom described the similarities between Archaeopteryx and Deinonychus. If dinosaurs were the new closest relatives of birds, then at least the most bird-like among them should be depicted with feathers (Ostrom himself disagreed with this though, and, according to Bakker, he fought the idea that Deinonychus should be feathered). Of course, which dinosaurs were most closely related to birds, and just how much of a gap remained between them and Archaeopteryx, allowed for considerable wiggle room and speculation.

A "carnavian" track maker
parachuting, from Ellenberger 1974.

In 1974, Ellenberger identified what he considered to be trace evidence for feathered dinosaurs from Lesotho, in the form of possible feather imprints and footprints which could have been made by small animals parachuting from trees, dragging their tail feathers in mud, etc. He illustrated a small bipedal dinosaur with feathers in the accompanying paper, along with totally speculative skeletal restoration of a new type of animal, nested within theropod dinosaurs, which he called "carnavians." Molnar, in 1985, dismissed the suggestion that these were feather imprints. Still, Ellenberger had become the first person to ever illustrate a non-avialan dinosaur, however hypothetical the species, with feathers.

Landry's influential 1975 restoration of "Syntarsus".

The next year, in 1975, a more famous feathered dinosaur illustration of a much better-known species was provided by Sarah B. Landry, drawn under the direction of Bob Bakker for his seminal article in Scientific American, "The Dinosaur Renaissance." Landry and Bakker depicted the small theropod "Syntarsus" (=Coelophysis) covered in overlapping feather-like scales or scale-like feathers, similar to Heilmann's "proavis", and a long tuft of feathers on the head. The choice of species was not a coincidence. Michael Raath, who had described Syntarsus in 1969 (the same year as Deinonychus), was quick to tout how bird-like it was in popular books and articles, and he suggested several times that it may have been feathered.

"Syntarsus" by Stout, 1976.

To understand the impact of this "first" feathered dinosaur, just look at the rest of the 1970s and early 1980s. It was Syntarsus, not Deinonychus, which was consistently drawn with feathers from then on. Many of these later reconstructions even directly aped Bakker and Landry's style of feather crest (or slightly modified it), making "Syntarsus with feather crest" a bona fide paleoart meme. Many of these left out the more subtle body feathers of the original, unfortunately, possibly influencing decades of "half-arsed" theropods with feather mohawks, up to including the dinosaurs in the 2001 film Jurassic Park III more than 20 years later. (One notable exception to the Syntarsus trend was also a watershed moment for paleoart. After reading Bakker's article, a paleoartist named Gregory S. Paul drew his first feathered dinosaur: an Allosaurus).

One of my favorite derivatives of Landry's Syntarsus illustration is one made in 1976 by William Stout and reproduced in Don Glut's 1982 edition of The New Dinosaur Dictionary. Though Glut pointed out that the feathers were speculative, they're probably less inaccurate than the legless snake it's eating! Also included in Glut's revised dictionary was one of the first illustrations of the theropod Kakuru, by Mark Hallett. Though just as speculative as Ellenberger's drawing (Kakuru is known only from two limb bones), it has more of a modern feel. The reclining theropod is decked out in long, filamentous feathers, rather than the broad, scale-like feathers of Landry's Syntarsus. It's worth noting that all of these early drawings of feathered dinosaurs had bare or scaly faces. This method of emphasizing the transitional character of early feathered theropods was probably inspired by traditional portrayals of Archaeopteryx, a "bird" with the head of a "reptile."

Feathered ornithischians, by Lorene Bjorklund, from
The Warm-Blooded Dinosaurs, 1979.

Many bird-like dinosaurs actually had feathers covering some or all of the face as well as the body, though some kind of "modular evolution" did occur to a degree in a few lineages (here's looking' at you, Darwinopterus). And some Mesozoic dinosaurs, most famously Kulindadromeus, seem to have feathers restricted to certain body segments with abrupt transitions between feathery and scaly regions. So, this much-maligned meme isn't necessarily out of the question, at least in early feathered lineages.

A few very early books featuring extremely prescient feathered dinosaurs came in 1978 and 1979. First, Julian May brought us perhaps the first renaissance-era dinosaur book for kids, The Warm-Blooded Dinosaurs, which featured not only a feathered Struthiomimus by Lorene Bjorklund on the cover, but several feathered ornithischians inside. An Iguanodon looks sort of ambiguously feathered (it might just be the crosshatched style), and interestingly, has a abrupt transition to a croc-scutes tail very similar to Kulindadromeus. There's also a skeletal of Microvenator with a fuzzy, Greg Paul style silhouette outline. The next year, Archosauria: A New Look at the Old Dinosaur by John McLoughlin includes some species with feathered faces for the first time, like his amazing, feathered Coelurus. It's odd as a modern reader to see that in both of these books, it's the more basal dinosaurs shown with feathers. Primitive ornithopods and "coelurosaurs" (considered a paraphyletic grade of early, ancestral dinosaurs at the time) like Syntarsus, Coelurus, and Saltopus are given feathers while deinonychosaurs like Saurornithoides and Deinonychus are not. The influence of the thecodont theory was still going strong, with birds thought to have evolved from the earliest dinosaurs rather than deinonychosaurs, which were exclusively known from the Cretaceous at that time.

Kakuru by Mark Hallett, from Glut 1982.

Ironically, Deinonychus, which out of all these dinosaurs was most similar to Archaeopteryx, did not become consistently or even frequently depicted with feathers until the at least the late 1980s. This was also probably due, in part, to artistic inertia. The original illustration of Deinonychus that accompanied Ostrom's description was so iconic that it was being copied well into the '90s. One of the first depictions of a feathered Deinonychus was a statuette produced by sculptor and founding father of Neopaganism Otter Zell (aka Oberon Zell). If anybody who owns a copy of this statuette wants to trade it for my right arm, please let me know.

After Syntarsus, the next non-avialan to become consistently depicted with feathers was Avimimus. From its first discovery, Avimimus was interpreted (and in some cases, like the supposed lack of a tail, misinterpreted) as being as birdlike or more than Archaeopteryx. Though commonly misreported online as having quill knobs, Avimimus actually had a flat ridge on the ulna which has been interpreted as a similar kind of support for the soft tissue of a feathered wing. Though not the same kind of direct evidence as quill knobs would be, most paleoartists ran with the suggestion, and most early Avimimus illustrations did portray it with feathers. John Sibbick restored it that way for David Norman's 1985 Illustrated Encyclopedia of Dinosaurs, and his version is a down-right throwback to the "pro-aves" of the early 1900s, with long, scale-like feathers on the outstretched arms and tail.

The first(?) feathered Deinonychus,
a statuette by Otter Zell, 1984.

1986 saw the publication of Bakker's The Dinosaur Heresies, filled with his own illustrations of feathered dinosaurs, including Deinonychus. (Stout and Paul had both continued to illustrate feathered dinosaurs up to this time, though not all were published or widely distributed at the time). Bakker's book opened up the door to more mainstream portrayals of this controversial subject, and it's no coincidence that many other artists were working the occasional feathered dinos into books and articles in the years that followed. Most notable among these were Paul's Predatory Dinosaurs of the World in 1988, and children's books inspired by it like The News About Dinosaurs. From then on, every "renaissance" era dinosaur book worth its salt included at least one speculative feathered theropod. Some books were bold enough to become the first to put "Feathers On Bloody Everything" (to quote the lament of some current-day feather detractors), theropods and ornithischians alike, as in 1988's speculative evolution book The New Dinosaurs by Dougal Dixon.

Two early restorations of Avimimus. Left: by Sergai Kurzanov, looking very bird-y.
Right: By John Sibbick from Norman's Encyclopedia, looking very much like Heilmann's "pro-avis."

By the time the first actual feathered dinosaur was announced in 1996, dinosaur fans, if not the general public, had been well prepared and had even come to expect such a find to be inevitable. The fact that this first find was Sinosauropteryx unfortunately reinforced some stereotypes that had been perpetuated by early renaissance era paleoart, such as short, fur-like feathers, naked legs and bellied, and half-feathered faces. Paul essentially invented the latter meme in an attempt to make his theropods look more bird like (by suggesting a sort of beak), and while this was his own personal speculation, many later artists ran with it, including in early drawings of Sinosauropteryx. The fossil appeared to support the hypothesis, which would have been an incredibly lucky guess for Paul if not for the fact that the snout past the eyes on that specimen had not even been fully prepared out of the rock (maybe, just maybe, because the scientists themselves were so influenced by Paul that they didn't expect any feathers to be found there!).

A modern feathered dinosaur:
 Tianyulong sculpture by Jason Brougham, 2016.

In a way, modern paleoart is still struggling to shed the memes and conventions generated by this pre-evidence feathered dinosaur artwork. Many artists are starting to critically examine just how far-off much of that early art really was, and to realize that it was, of course, just pure speculation about what may be, not informed speculation based on fossil evidence. Many dinosaurs, especially non-avialan coelurosaurs, had a feather covering that was much more Archaeopteryx-like than any of those pioneering artists could imagine. It's only recently that many of the best artists have been going back for inspiration not to Landry and Paul and Bakker, but to the earlier artists who were drawing Archaeopteryx and the various "pro-aves". This new era of paleoart, part of what Tom Holtz has dubbed the "Dinosaur Enlightenment," will no doubt generate its own set of tropes which will need to be overcome by the next generation of artists. We're already seeing that with the dogmatic insistence from some quarters on a certain standard of feathering for all dinosaurs when the fossils are starting to suggest much more variety. With any luck, the internet will allow this new generation to push forward and more widely and rapidly share new ideas and possibilities about what feathered dinosaurs can be.

You're Doing It Wrong: Protobird Toys Edition

The new Carnegie Velociraptor figure.

I'm pretty slack when it comes to keeping up with my Twitter account, but if one thing can get me to break my one post per month general rule, it's "somebody is wrong on the Internet!" Or, in this case, "somebody made an inaccurate dinosaur model under the auspices of museum-approved accuracy!"

Let me preface this post with a few disclaimers. One, I have no problem with people making dinosaur art however they like. But I feel it's very important to draw a distinction between dinosaur art in general and paleoart. Creating a drawing, painting, or sculpture under the guise of paleoart implies that some degree of research went into the piece. When evaluating paleoart, critics are completely entitled to demand as rigorous an approach to accuracy as the evidence allows. Of course, speculation must be included to some degree, but the baseline expectation is always that basic facts and plausible inference will be taken into consideration.

Dinosaur art is a for of pop art, a purely artistic expression. Paleoart is a representation of a scientific hypothesis about the life appearance and behavior of an extinct organism.

This recent Twitter kerfuffle was due specifically to the debut of a new dinosaur figure in the Carnegie Collection produced by Safari Ltd. I've been a big fan of the Carnegie figures (and the sometimes nicer quality Wild Safari sister series) since I was a wee lad buying my first Carnegie Pteranodon in a little hobby shop for about $2, and pining over their massive Brachiosaurus. Carnegie figures have obvious appeal for scientifically-minded young dinosaur fans. First, most of them are made with a consistent scale (usually 1:40), so when you line them all up you can see how big each animal was compared to one another. Second, they are marketed as having the highest level of scientific authority: the Carnegie Museum stamp and assurances that hey are approved by actual paleontologists gives them the weight of authority and accuracy not found in many other toy lines. Sure, many of the older figures, including my Pteranodon, are now sorely out of date, but this is due largely to the Science Marches On effect rather than a basic, original inaccuracy.

However, like many paleoartists, the sculptors at Carnegie and related accuracy-minded/marketed toy lines seem to be having, well, let's say a little trouble adapting to the feathered revolution. Trained in the '80s and '90s drawing and sculpting Paulian, reptilian dinosaurs, it's been a steep learning curve for many of these artists to switch to bird-like, feathered dinosaurs, which many artists don't even realize requires a crash course in avian anatomy, rather than reptile or mammal anatomy, to get right. This is why expert consultants are so important in these projects--artists simply need help catching up with the latest research. Unfortunately, they're usually getting bad advice from "experts" who simply do not care.

The newest entry in the Carnegie Collection is the old favorite Velociraptor. Carnegie had previously released a scaly Velociraptor, but like many of their other models, they have commendably updated it to try and reflect current science. Unfortunately, it seems they mainly achieved this by slapping feathers over the basic original model with no regard for the fact that feathers inherently change the entire presentation and outline of an animal. Unlike fur or "protofeathers", which just fluff up the outline of an animal, feathers are more like a mobile exoskeleton that re-defines the entire body.

My own hypothesis, based on fossil and phylogenetic evidence as well as
inference from living analogues, about the life appearance of Velociraptor.

That being said, the Carnegie Velociraptor is not that bad overall and is very nicely sculpted, with an interesting and plausible color scheme. The feathers look fur-like, true, but this is not unprecedented among different lineages of large ground birds, and so is not unlikely in flightless protobirds (though they'd probably still be longer). The main problem here is the wing. Yes, wing. Like chickens and ostriches, protobirds possessed fully-fledged wings despite being flightless or nearly so, and retaining large claws on the fingers (yes, chickens, ostriches, and many other birds have hand claws--they are not some prehistoric protobird relic!).

Specifically, the wings in the new Velociraptor figure are very small, with short feathers, and are present only on the forearm, not the hand, making them only half a wing--literally, since they're issuing the primary feathers. We do not have direct evidence of primaries, but we have never found a single example of a maniraptoran that has secondaries but not primaries, and so it should be assumed they were there by default. As for secondaries, we have direct evidence in the form of quill knobs for this species. Quill knobs are not found in all feathered animals, let alone all flying birds, and seem to be associated with strong attachment either due to high-stresses during flight or other flapping behavior and/or especially large/long individual feathers.

Long story short: Not only did Velociraptor have wings, it probably had larger wings than many other dromaeosaurids. AFAIK not even Microraptor and Archaeopteryx had quill knobs to support their wings.

The new Carnegie figure, on the other hand, barely has wings at all. What went wrong?

Some insight can be gained by a recent series of incidents involving sculptor Dan LoRusso on the message board of the Dinosaur Toy Blog. LoRusso Is an amazing artist and is responsible for some of my all-time favorite dinosaur figures released in the Boston Museum of Science Collection by Battat (now being re-released with updates and new figures under the Terra brand). Collectors were understandably excited about the fact that the Battat series was coming back after nearly two decades, though the excitement was dampened a little by some obvious accuracy issues in the new figures, specifically when it came to the feathered species (or species that should have been feathered).

LoRusso was criticized online for producing a very well done but very inaccurate therizinosaur figure which completely lacked feathers. It would have fit right in with the excellent quality and Paulian style of the original series... back in 1994. But in 2014, when we have incontrovertible proof that therizinosaurs were not only feathered but that at least smaller species were very densely feathered, it is simply bizarre to see a featherless figure in a line that is being marketed as scientifically accurate. LoRusso stated that his consultants told him larger therizinosaurs would have been featherless. It's not LoRusso's fault that he somehow was able to sculpt a bald maniraptoran in the year 2013. Somebody who did not know what they were talking about and claiming to be an expert in paleoart just because they work in the related field of paleontology told him to do it, and he very reasonably believed them because they were an "expert", though obviously they had misrepresented themselves.

And there's the biggest problem and probably the answer to the question of what is going on with these strange and obvious inaccuracies. Paleoart consultants for major projects tend to be, often but not always, simply terrible. They seem not to know what they are talking about. Not only that, but second-hand reports suggest that they often simply do not care. Seriously: When asked about blatant inaccuracies creeping into paleoart-based projects like toys or books or even press releases, at least one anonymous paid paleontological consultant stated that they don't care what dinosaurs looked like in life, and so would presumably rubber-stamp any abomination that came across their desk.

@mpmartyniuk You know I've spoken to working palaeontologists who even say that they >don't care< about depicting life appearance?
— Darren Naish (@TetZoo) October 4, 2014

Here's a prime, objective example: The Carnegie Collection Caudipteryx zoui figure. This species was first found in 1998, a complete skeleton with feather impressions. There are plenty of photos of Caudipteryx fossils that show crystal clear how the feathers attach and were ignored completely for this figure. The artist might have copied some inaccurate depiction rather than doing actual research or glancing at a fossil, and the consultant approved it because they didn't know or didn't care about the relevant details. The Carnegie Caudipteryx proves that consultants are utterly useless and often have no clue what they're talking about. Any one of us can compare the model with the fossil and show that the model is objectively wrong in major details. Look for yourself:

Caudipteryx wing fossil: note the primaries, longer than the hand, anchored along the second finger,
and lack of a clawed third finger.

Carnegie Collection Caudipteryx figure. Note the wing feathers anchored everywhere along the arm
EXCEPT on the second finger where they belong, and the incorrect number of fingers.

The wings/arms of this figure are wrong in just about every single way possible, and these are not minor details. Yet it's marketed as accurate and "paleontologist approved" in a series bearing the name of a major scientific institution!

It's not fair to ask all working paleontologists to know or care how their research into fossils translates into life appearance. Matching osteological and behavioral correlates with the structure and anatomy of living analogues could almost be considered a distinct field separate from actual paleontology. This is something paleoartists can and do think about and research constantly, but would almost never need to enter into the research when describing fossils. There's really no reason for working paleontologists to keep up to date with developments and research that go into paleoart.

The solution? Don't ask these paleontologists to consult! Just because someone is a paleontologist does not make them an expert on the life appearance of any given species of prehistoric organism. The examples cited above were, allegedly, all approved by "expert" consultants. This simply proves the consultants that are being employed are utterly failing at their job. I hate to say it, but there are legions of paleoartists and other dinosaur fans online who jump at the chance to criticize and nitpick and otherwise consult on these things for free. It's just that by the time the product is released, it's too late to do anything about inaccuracies. If companies that use paleoart would simply post concept art beforehand on, say, Facebook, they could probably get much better advice for free. Or, preferably, they could employ actual paleoartists as experts, hopefully artists who specialize in researching the life appearance of a given subject group of organisms.

Darren Naish, Mark Witton, and John Conway recently published an article on the (shameful) state of  paleoart as used in professional and commercial contexts. Companies like Safari Ltd. would do well to listen to their advice.

People Think Feathered Dinosaurs Don't Look Scary. They're Right.

This short article on io9 pretty well encapsulates an area of frustration for artists and scientists in the age of feathered theropods.

Ooh, I'm shakin' in my boots.
Photo by Simon Rumble, licensed.

The implication is, right from the title, that it's common knowledge most depictions of feathered T. rex are not cool. Feathered theropods are widely derided by the public because feathers make these scary reptilian monsters less scary. In a recent Facebook discussion, I took one of those "short pelt raptor" images to task for inaccuracy (you know, the kind that pays lip service to the idea of feathered theropods, but with the minimum possible change to the classic silhouette, with a cat-like short pelt rather than a bird-like poof of feathers engulfing the body). In my reply I kind of hypothesized that there's an evolutionary psychology* reason for our aversion to feathered theropods and our cat-like concessions to the idea.

As Andrea Cau has pointed out, paleoartists (myself included), consciously or not, often employ all kinds of subtle tricks to make feathered theropods look "cool". Leaving the face scaly and reptilian is a popular trick; his body might say "Big Bird", but his face tells you he means business. Face fully feathered? Introduce an eagle-like lowered "brow" or some kind of eyebrow analogue so his facial expression can look "mean". Make sure his mouth is open or he's prominently displaying his other weapons in a ninja-like fighting stance. And be sure if you make him colorful, use high contrast, red and black if possible, and light it so his face is in shadow--that way you know he's thinking evil thoughts. This might also allow you to add eye shine making the eyes look like demonic embers! (check back to the io9 article and see how many of these points that T. rex hits).

In the comments to the io9 article, there were the predictable bouts of resistance to the idea that T. rex could have had feathers at all. "It was too large! Large mammals don't have so much fur in hot climates!" The problem with comparisons to large mammals is that feathers are very different in structure from fur, and have very different insulating properties. Fur is mainly used to keep an animal warm, but thanks to the fact that feathers grown in adjustable, planar layers, and are better at trapping and regulating air flow, many large birds use their feathers to very effectively keep themselves cool by circulation while blocking the skin from absorbing direct sun. It may actually have been disadvantageous for a large animal to lose its feathers, especially if it lived in a hot sunny climate. The fully-feathered Yutyrannus was not significantly smaller than any but the largest T. rex. Most T. rex specimens fell short of the 6.8 tons estimated for the most gargantuan known adults like Sue, that is certainly not the species average size!

But, there was one comment that played right into my ego-psych hypothesis. The commenter basically stated that we know juvenile T. rex had feathers, but there's no reason to think adults kept them. Except even that premise is wrong. There is in fact zero direct evidence to support the hypothesis that T. rex juveniles had feathers, let alone that they had them and then lost them. It's simply easier for people to assume that a baby animal, which is supposed to be cute, had feathers, which we psychologically associate with cute animals. 

Are one of these things is not as scary as the other?
Illustrations by M. Martyniuk, all rights reserved.

It is actually less of a stretch (i.e. more parsimonious) to hypothesize that based on its phylogenetic bracket, T. rex had feathers and retained them throughout its life, than the hypothesis that T. rex was born with feathers, lost them because they became disadvantageous at some unspecified weight, then through some unknown developmental pathway replaced its feathers with the kind of thick, scaly skin it is usually depicted with and would need to protect itself from the sun/injury if it lacked feathers. But this convoluted thinking is easier for people to accept because T. rex is the king of all monsters, and monsters are by definition not cute.**

The sad fact is, T. rex may not have looked all that cool. I think John Conway and others have brought this up before. It, and many if not most other dinosaurs, may very well have looked really, really stupid to us. Nature doesn't care if an animal looks intimidating to a species that evolved 66 million years later in a completely different environmental context alongside a vastly different set of predators. Our brains are programmed to find mammalian and reptilian predators scary at least in part* because we evolved alongside these and our survival depended on it. We had no such pressure for most kinds of birds***, and maybe coincidentally, we find very few kinds of birds the least bit intimidating. We have to be told/shown that a cassowary is even capable of being dangerous, and people still constantly trot this out as a surprising fact, despite the fact that it has very few physical differences from a Velociraptor, other than being much larger! 

So, if your average Joe met a non-avialan theropod in real life, the reaction might be less like any of the raptor scenes in the original Jurassic Park and more like Newman vs. the cute, colorful, silly, hopping (read: bird-like) dilophosaur - bemusement leading to injury.

* I know evo psych is mostly made up of untestable just-so-storys. But it's still fun to think about.

** That's sarcasm. Tyrannosaurs were not monsters, they were plain old regular animals. A lesson people forget from the original Jurassic Park (probably because they're not actually depicted hat way in the movie, despite the fact that the characters talk about it).

***Raptors seem to be the exception. Probably because they preyed on early humans, and maybe also because of their mean-looking "eyebrows"?

What Does T. rex Say?

"Hissssssssssssssssss!"
T. rex holotype specimen. Photo by Scott Robert Anseimo, CC BY-SA 3.0.

It's an iconic scene in every dinosaur movie: the huge, conquering carnivorous theropod rears back and lets out a terrifying bellow. Sound effects artists spend huge amounts of time sampling vocalizations from various animals to create just the right mix to create an unfamiliar, otherworldly roar. And, of course, everybody knows that pterodactyls let out harsh, echoing, prehistoric sounding screeches.

But how close to reality are these sounds? Do we have any ways of using science to figure out what dinosaurs and other stem-birds may have sounded like? Do we have evidence that they made sounds at all?

Review: Papo Archaeopteryx

Like many paleontology fans, I have a pretty big collection of little plastic dinosaur toys. Most of these I got when I was a kid and have held onto since, but every so often a nice looking model is released that is too cool to pass up. This new Archaeopteryx figurine from Papo was one of them.

Oh, Hi, Bohaiornithids!

It's not often that we are introduced to a large new clade of stem-birds*, but a new paper by Wang et al. finds support for just such a thing among the enantiornithes. Named Bohaiornithidae, the family unites a few previously-known similar-looking opposite birds with two brand new species.

Phylogeny of Bohaiornithidae, modified after Wang et al. 2014.

Iridescence in Simple Feathers - The Case of the Blue Troodon

Iridescent feathers in troodontids - possible? Image by Matt Martyniuk, all rights reserved.
(Supreme dino fans may recognize the pose even from this small clip...)

I've written a lot about the various ways feathers get their color in order to create some rough guidelines for paleoartists restoring feathered stem-birds. I recently had a quick discussion over at DeviantArt with one of the best currently-working paleoartists around, Emily Willoughby, over the plausibility of a blue Troodon. Note that it has a few other anatomical issues that make it somewhat less than a fully accurate rendition, but is the coloration one of them? I wasn't sure, so I did a little extra digging to find out. As usual, this research is cursory and I welcome any additional input on research or details I may have overlooked.

You're Doing It Wrong : Dino Foot Scales

Above: Our subject matter.

It's often said by those who support a strict phylogenetics-based system of naming life that it's only by restricting well-known names from neontology (the study of modern organisms) to crown groups can we avoid making unjustified assumptions about members of stem-groups.

These kinds of unjustified assumptions have been rampant in the history of studying stem-birds. Archaeopteryx has traditionally been depicted, incorrectly, with a reversed hallux, and occasionally even with beak-like structures, simply because it's a "bird", and those are features all birds have. Except Archaeopteryx is not a true "bird", it's a stem-bird, more closely related to birds than to any other living animal group, but not a member of the group that includes all modern birds. It's fair to assume that an extinct member of the duck lineage, like Vegavis, had a bill, but that's not necessarily so for, say, Patagopteryx, despite the fact that it is usually referred to as a "bird".

Modern bird feet, by Philip Henry Gosse, 1849, public domain. Note overlapping scutes on
the top surfaces, and pebbly, polygonal reticulae on the bottom surfaces.

Most paleoartists have absorbed these kinds of warnings, and do a good job of avoiding obvious errors based on typology, the assumption that all species in a certain "type" share "key characteristics." But there are some typological memes in the bird lineage that are more pernicious, possibly because their actual evolution is something most artists don't think about very much.

Take, for example, the bird-like scutes that are almost universally illustrated covering the tarsus (upper foot/lower hind limb) of dinosaurs. Is there any evidence that these were actually present in any given group of non-theropod stem birds? Well... no. Not that I'm aware of (if you know differently, please comment!).

Sinosauropteryx prima with tarsal scutes.
Image by Matt Martyniuk,  licensed.

I'm not sure when this meme began, and if it's related to the Dinosaur Renaissance when the link between birds and dinosaurs was re-established. Looking at some Charles Knight paintings, such as his famous "Leaping Lealaps", it appears that the feet of his theropods were scaled based on modern lizards (more on the differences between lizard scales and other types of "scales" below). Bakker's influential early restoration of Deinonychus does not include any obvious scutes on the feet or tarsus. Mark Hallet, on the other hand, did include what look like oblong bird-like scutes on his theropods. At any rate, it's hard to deny that "bird feet" are typical of almost all modern reconstructions of dinosaurs, including my own, and are not limited to theropods. Bird-feet are often restored on ornithischians and even pterosaurs.

Of course, like many paleo-memes that developed during the 1980s, the main idea seems to be using this as a flourish to make otherwise scaly dinosaurs seem more bird-like. And thanks to skin impressions, we know that many dinosaurs had scales, right?

Follow-Up: Judith River Formation = Oldman Formation

In a previous post, I hung my tentative re-identification of the holotype teeth of Deinodon horridus on a rough correlation between the Judith River and Oldman formations, the latter of which is more precisely dated and, more importantly, contains Daspletosaurus torosus, which is a candidate for the owner of Deinodon teeth.

While researching a different topic, I stumbled across a more definitive published correlation of these two formations I wasn't previously aware of. In their 2001 paper on the stratigraphy of the Two Medicine Formation, Horner et al. discuss the correlation of parts of that formation with the Judith River. Horner et al. note that the Judith River can be separated into two basic units divided by a disconformity, corresponding with a marine transgression (when the terrestrial ecosystem was swamped by the rising of the Western Interior Seaway, the sediments deposited by which appear to have been lost in this instance).

Helpfully, Horner et al. note that it is from the lower unit that Hay collected numerous dinosaur teeth which were later described by Leidy as the infamous tooth taxa such as Deinodon, Aublysodon, Trachodon, and Troodon. More helpful still, the paper provides a handy chart showing the arrangement of the strata and including points at which radiometric dates have been taken. The base of the lower Deinodon-bearing unit is dated at about 78 million years old. The next available date is from just above the disconformity (i.e. after the seaway had retreated again) and shows an age of 75.4 million years ago. That's narrowing it down, but there's no date from within the formation from just below the disconformity, which would give us an upper boundary for the Deinodon strata.

But, there's hope. Horner et al. note that Rogers (1998) suggested the disconformity itself probably correlates to around the Willow Creek Anticline in the middle Two Medicine Formation (which contains the famous Egg Mountain Maiasaura nesting site). This segment of the TMF has been dated to 76.7 Ma ago, which may give us a rough upper boundary for the age of Hay's fossil tooth collection.

So, based on this paper at least, it looks like Deinodon and friends were collected from rocks aged somewhere between 78 and 76.7 million years old. Which is about the same age range as the Oldman Formation to the north. So, Deinodon horridus and Daspletosaurus torosus did indeed live at about the same time and in the same region (there were no checkpoints at the US-Canadian border back then!), making it more likely that they represent the same species, and the possibility that Deinodon actually represents Gorgosaurus less likely.

Looks like I'm going to have to create a new tag for Arcane Biostratigraphy and Geology Stuff...

Oh, and somebody in the comments last time asked me to get into Trachodon. This is definitely a subject for a longer blog post, though I'm a bit less excited about it because I'm more pessimistic that it's identity is knowable. But maybe this new info can help us get started. I already mentioned that the Trachodon teeth appear to come from the same strata as Maiasaura (and it's well known that Trachodon's contemporary Troodon formosus is reported from Egg Mountain as well, though T. formosus is also reported from pretty much everywhere and everywhen else...). Could Maiasaura be Trachodon? Perhaps! But it could also be Brachylophosaurus, or maybe even Gryposaurus. And... there's been a rumor going around for a while now that Trachodon teeth are referable to Lambeosaurinae. Which lambeosauines are known from this time and place that could fit the bill? Both Parasaurolophus and Corythosaurus have been reported from the uppermost Oldman, though these may be too young. Hypacrosaurus sp. seems to have been contemporary with Maiasaura, so that could be it...

Yeah, you can see why I'm pessimistic. Tyrannosaurids are rare, and there tend to be only one or two species of tyrannosaurids present in any given ecosystem. Hadrosaurids are... the opposite.

References

Horner, J. R., Schmitt, J. G., Jackson, F., & Hanna, R. (2001). Bones and rocks of the Upper Cretaceous Two Medicine-Judith River clastic wedge complex, Montana. In Field trip guidebook, Society of Vertebrate Paleontology 61st Annual Meeting: Mesozoic and Cenozoic Paleontology in the Western Plains and Rocky Mountains. Museum of the Rockies Occasional Paper (Vol. 3, pp. 3-14).

Did Jurassic Park Name T. rex?

T. rex illustration by Matt Martyniuk, licensed.

There's a pretty interesting historical paleontology thread happening over at the Hell Creek forum. In the most recent issue of Prehistoric Times, one article claims that the influence of the film Jurassic Park, released in 1993, included popularizing terms like the name "raptor" for dromaeosaurs (unquestionable) as well as the abbreviation T. rex for Tyrannosaurus rex. Did JP really give the world "T. rex"?

Deinodon's Identity Revisited

My recent illustration of a specimen traditionally assigned to Gorgosaurus was labelled Deinodon, usually considered a likely synonym. But is it really? Deinodon is, in fact, much more likely to have been Daspletosaurus all along. Read on for the nitty-gritty stratigraphy!

You're Doing It Wrong: CGI Feathered Dinosaurs

YIKES.
Promotional still from When Dinosaur Roamed America,
Discovery Channel 2001

CGI dinosaurs have come a long way since Jurassic Park. Unfortunately, due to the discovery that these animals were far more bird-like than reptile-like, that direction is straight down a seemingly bottomless pit. For many CGI artists and animators, the key concept that is presenting a very steep learning curve is the idea that these animals need to be restored as feathered dinosaurs, not "dinosaurs with feathers." It looks like many people modeling feathered dinosaurs start out by rendering a featherless dinosaur and then using various modeling or rendering tricks to add the feathers. As you'll see below, this never, ever works.

Supporting the Dinosaur/Bird Link in the Era of the MANIAC

I'm posting this more as an open question than a statement of my own opinion, so comments appreciated!

A new paper out in Paleobiology by Verracchio et al. describes the porosity of Troodon formosus eggs and uses the data as evidence to support the hypothesis that troodontids brooded their eggs, like modern birds and other known maniraptorans, rather than burying them, like crocodilians and some other modern birds. In and of itself, this conclusion is interesting in that it pretty much solidifies nest brooding (as opposed to burial) as the ancestral trait for modern birds, and for maniraptorans (or at least chuniaoans*) in general.

Study coauthor Darla Zelenitsky with Troodon formosus nest. Photo by Jay Im, University of Calgary.

I hate to admit it, but my first thought when reading the headline of this news article from PhysOrg was that, yeah, we all assumed that anyway. The unspoken "rule of cool" is that science tends to be more exciting when we find evidence that contradicts previously well-supported hypotheses, rather than confirming hypotheses we all took for granted. Sure, finding the Higgs-Boson was exciting, but not nearly as exciting as not finding it, which could have led to new physics. I assumed, and I'm sure many others did as well, that troodontids brooded their eggs, based on the reasonably secure hypothesis that oviraptorids (which are known to have done so) are more basal. This behavior in troodontids was even depicted nicely in 2011's Dinosaur Revolution. So score another one for phylogenetic bracketing!

(Of course, this is not to imply that all chuniaoans must have brooded their eggs. It's entirely possible that reversals to burial nesting occurred, as with modern megapodes, and this seems especially likely for very large species like some dromaeosaurines. But the odds that any given chuniaoan would not be a brooder are low.)

Arctic troodontids, anatomy based on Troodon formosus.
Matt Martyniuk, all rights reserved.

Aside from all that, the assertion in the PhysOrg headline struck me as particularly meaningless. How could a study of troodontid brooding lend support to the dinosaur/bird hypothesis? This statement would have been accurate a decade ago, but not today. The reason is the moving goalposts of the dinosaur/bird opposition.

The Flying Longipterygids

I started to get a little carried away in my previous post on longirostrisavisine tails, so I decided to split the finished post in half and use this one to delve a little more into the possible implications the odd tails and clawless wings of that group would have had on their flight styles.

As mentioned previously,  longirostravisines, unlike most other enantiornitheans, lacked claws on all three of their manual digits, a trait apparently acquired separately of the Carinatae (Ichthyornis + modern birds). In the last post I speculated that this might have reflected a distinct flying (or at least landing) style. But how much do we really know about longipterygid flight?

Fig. 10 from Close & Rayfield 2012, click to big it up. Note the three general clusters of continuous-flapping (red), soaring and flightless (yellow and gray), and bizarro enantiornithean style (green).

In their recent paper, Close & Rayfield attempted to correlate wishbone (furcula) shape with flight style. Many enantiornitheans in the analysis plotted together in a cluster well separated from modern birds, but close to basal avialans like Confuciusornis which are usually presumed to have been gliders. Oddly, in the Close & Rayfield figure reproduced above, Rapaxavis (no. 103), a longirostravisine, clusters with the weirdo enantiornithean-style fliers, while more basal longipterygid Longipteryx (no. 98) groups with continuous-flapping fliers and is marked as an ornithurine (something seems off here).

In another recent open-access paper on Mesozoic bird flight styles, Wang et. al (2011), using wing element and primary feather length ratios, found Longirostrisavis to be a continuous-flapping flier, and Longipteryx either a continuous-flapping or a flap-gliding flier, roughly the opposite result of the Close & Rayfield furcula-based analysis.

 The fact that two independent analyses using two distinct methods produced different results probably does not bode well for the precision of either method. Clearly, we need to figure out better methods to more reliably tests flight styles in extinct birds.

Did Sinosauropteryx Have "Protofeathers"?

Life restoration of the S. prima type specimen by M. Martyniuk

Over at Jaime Headden's blog The Bite Stuff, Jaime recently wrote a great article about the newly discovered basal coelurosaurian or orionidian Sciurumimus. The article touched on the fact that its feathers were reported as stage 1 (simple, unbranched filaments, often referred to as "protofeathers") in the parlance of feather development researcher Richard Prum. In the comments, Heinrich Mallison pointed out that they *look* like stage 1, but as Foth showed, crushed feathers (even those of modern birds) often look much more primitive than they are due to taphonomic effects. (I've mentioned before in a few places that it's really unfortunate Foth's important paper seems to have been mostly ignored by others writing about fossil feather types). Jaime defended this by saying that Sinosauropteryx, which is more derived, also is 'generally agreed' to have had stage 1 feathers. It's true that this is the general agreement in the literature, and it's also true that the general agreement has been challenged (effectively, in my opinion). I posted a reply to the article, but the fact that this keeps coming up lately has made me think that maybe I should be trying to publicize this more widely, so my comment is reproduced below along with some more commentary on the issue.

The Strange Bird Dalianraptor cuhe

Type specimen of D. cuhe, originally posted by Andrea Cau

I'm interrupting my regularly scheduled upcoming blog posts to bring some attention to a little-known Jehol bird: the strange 'jeholornithid'-grade species Dalianraptor cuhe.

D. cuhe has spent the last several years as a species in obscurity, even among most paleontology enthusiasts. I recall my first glimpse of the type specimen, wondering over the seemingly-complete remains of an "undescribed possible dromaeosauird" in a low-res photo posted online in the early '00s. I can remember saving the image to my reference folder, hoping that one day I'd be able to update the file name. It's a fascinating animal, but... is it real?

The Tail of Shanweiniao

Fossil tail feathers of S. cooperorum, from O'Connor et al., 2009.

As a follow-up to last month's post on the smallest Mesozoic theropods, here are a few additional observations on the small longipterygid Shanweiniao. Like other longirostravisines*, Shanweiniao cooperorum had reduced "hands" entirely lacking claws. This reduction of wing claws seems to have occurred independently of modern birds within this uniquely specialized group of enantiornitheans. (Euornitheans seem to have lost the bulk of their wing claws around the level of Carinatae, though many modern birds still retain at least keratinous claws on their wings, and longirostravisines may have as well).

S. cooperorum itself is most well-known for its elaborate tail made up of six ribbon-like feathers. Those feathers overlapped at the base, and may have acted as an air brake for precise landings with the feet on small branches. It's possible that most other enantiornitheans, which lacked long feathery tails and also retained wing claws, landed by simply smacking clumsily into tree trunks or brush and grabbing on with all four limbs.