Revisiting the Fisher King


I know, I know, my last post promised a series on reconstructing Acrocanthosaurus in multiple views - bear with me, as this is actually part of that series. Remember that both animals have stuff sticking up on their backs, so I want to be able to compare and contrast those elongated neural spines...and how those differences should impact reconstructions of the animals. But to do that I had to update this skeletal, as new information had rendered the older one no longer tenable.

Besides, Spinosaurus is cool! For one, it's the only dinosaur in the Jurassic Park series to tangle with a T. rex and emerge victorious (no matter how unlikely that outcome was). It's probably the longest theropod we know of, and may have been the heaviest as well. Yet counter-intuitively it shows specialization for piscivory (fish-eating)...maybe in JP3 the spinosaur mistook the T. rex for a really large lung-fish?

Tongue firmly out of cheek now, Spinosaurus has lit up imaginations partially due to its size, but also because there was so much you had to imagine to try and reconstruct the animal. Until the last decade or two it was sort of a theropod Rorschach test where you could project any sort of oversized monster theropod onto its scant (and now lost) remains. This brings a thrilling "Sherlock Holmes" quality when trying to imagine the living animal, but for most of the last century serious attempts to reconstruct Spinosaurus have been more frustrating than titillating.

Darren Naish has an excellent write up of the history (and tragedy) of of the type specimen of Spinosaurus, which I won't duplicate here. The long and short of it is that WWII claimed the fossils as another victim of the conflict. The already-meager remains lost, paleontologists were stuck with the original description and some somewhat uninspired sketches as the only link to the past.

A series of fortunate events occurred in the latter half of the 20th century that allowed for a more accurate interpretation of Spinosaurus to emerge. For one, other spinosaurids were found. Baryonyx from the U.K., and Nigerian Suchomimus, started to paint a more complete picture of what these animals were like. They had bizarrely long snouts that seemed to resemble a gharial as much as a traditional theropod. Suchomimus even had a smaller version of the enlarged neural spines on the back:


The amusingly-named Irritator from South America further clarified the relationships and anatomy of spinosaurids. But the real breakthrough was the re-discovery of several photographic plates of the original material. While Spinosaurus wasn't the most complete specimen, having photographs at least made it possible to ensure that what was found is incorporated accurately into a reconstruction.

Among other details, the image also shows what had been the basis of attempts to restore the shape of the elongate sail or hump on the back: Stromer's original interpretation for the position of the elongated neural spines. In particular, notice that the tallest one is set directly in front of the sacrum here, while the only associated tail vertebra (at the far left of the picture) has a very short spine. That has lead most people to infer that the spine started quickly after the neck, grew to ridiculous heights over the pelvis, and then quickly dropped off again. Indeed, this is the interpretation that I used in my first attempt, and has been widely seen in such disparate and reputable scientific endeavors as Jurassic Park 3, the Carnegie Collection of "museum quality replicas", and Greg Paul's reconstruction in his Princeton Field Guide to Dinosaurs.

And they're in excellent company (whereby I arbitrarily define myself as "excellent company"). I had been concerned with Stromer's original interpretation for the placement of the tallest neural spinse - no vertebral body (centrum) was preserved, but the change in the angle of the spine seemed pretty extreme compared to the previous dorsals, especially right in front of the sacrum. My solution was to assume it was a sacral neural spine. This largely preserved the traditional appearance of the "sail", but provided a bit of breathing room for the change in orientation.


Luckily for us, Andre Cau and Jamie Headden were busy mulling over this specific issue, and came to a much more likely conclusion, that the backward-oriented neural spine was actually an anterior caudal. Looking at a host of dinosaurs with elongate neural spines, they noted that in general you never seen backward-canted spines in front of the hips, you always see them after it. There is a bit more detail to the argument (which I encourage you to read on their blogs), but in essence they make a very compelling case.

And so it was back to the virtual drawing board. I made some other corrections from my previous attempt - there had been some scaling issues with the neck vertebrae that had given my reconstruction a thinner Baryonyx-like profile in the neck. Also, it appears that the necks of these animal don't have as much of the traditional theropod S-curve, so that was changed as well (although I still don't buy the extreme hang-dog look that Greg Paul has started to restore his spinosaurs with). The results are a stockier animal, with a more elongate sail (or hump):


Looking at the rigorous reconstruction, it's clear that there's still quite a bit of uncertainty in the skeleton, although not all of the missing parts are created equal. Much of the pelvic girdle is known from Irritator, as is the back of the skull. Also, some unpublished specimens shed light on this, even if they aren't documented well enough to be official parts of the reconstruction. Still, there's a bit of ambiguity about the exact limb proportions, the length of the tail, and the exact shape of the sail.

Speaking of which, how should those tall neural spines be restored by artists doing life reconstructions? Is it a sail, was it supporting a hump of tissue like a bison, or was it simply a muscular ridge? We'll get back to that subject in a bit, after looking at Acrocanthosaurus.

Until then, best wishes to one and all for a wonderful 2012!

Falcarius: bizarre sickle-cutter


The truly strange looking animal above is Falcarius utahensis. It's an early, omnivorous member of the theropod clade known as therizinosaurs. Not only does it look weird, it's also a bit different from other skeletals you may have seen on the web. Join me after the break for a bit of a discussion about Falcarius, and the challenges I faced with this reconstruction.

I should warn you, this won't be a tutorial on how I make my skeletal reconstructions. That would certainly be a fun series, but it would require quite a lot of time to do properly, so for now it'll have to wait. But there are still several points worthy of discussion.

First off, the animal was discovered in a bone bed of disarticulated individuals. The good news is that most of the individual elements are known, but the down side is the bones aren't all from the same sized animals. That means that cross-scaling is needed to restore the skeleton, but even that presents a challenge; the usual method of cross-scaling involves double-checking the results against the proportions of close relatives. Alas, in this case the fossil record for the base of the therizinosaur family tree isn't well known, and what is known makes it clear that Falcarius has very different proportions than it's closest known relative: Beipiaosaurus.

(Image copyright Greg Paul)

When the original description of Falcarius was published in 2005, it came with the skeletal drawing at left. Obviously I don't agree with those proportions now, but at the time it had been done when fewer bones had been excavated, prepared, and described in detail, so Greg Paul had to try and scale them based on a smaller amount of material to compare with.

In fact, given the difficulty of restoring the proportions I intentionally avoided doing a Falcarius skeletal reconstruction for several years. I might have avoided it all together, but towards the end of my tenure at the WDC we mounted a cast of Falcarius that Gaston Design produced. Working on that skeleton I was able to not only measure and photograph all of the elements, but spend time looking at how the individual elements were matched up. Some parts of the cast's vertebral column are from different sized individuals (an unavoidable consequence of trying to piece together a skeleton from several different individuals). In other cases, vertebrae I had assumed to be from different sized animals were in fact crushed.

In addition to the hands-on data, Lindsay Zanno had been hard at work publishing more detailed information on Falcarius (this is actually notable, as not all researchers are as timely with getting more detailed descriptions of a new animal into print).

As the information piled up I felt that a skeletal was possible to be done. I still didn't tackle it though, as there were plenty of "low-hanging fruit" skeletals that could be done from less-challenging animals.

As luck would have it, I ended up being asked to produce a skeletal of Falcarius for a display in the new Utah Museum of Natural History building (side note: the new UMNH building just opened, and houses one of the most impressive natural history displays in North America, go see it!).

Since I was working with the UMNH, I got valuable input from several of the researchers who worked on the specimens. They were able to provide additional information - I won't go into the nitty-gritty of it (although you may if you would like), but I wanted to point out that the end result was quite a surprise to me. And little is more satisfying than when you are really surprised at the end of a skeletal reconstruction.

Resulting skeletal in hand, you can compare it to the most recent studies of the therizinosaur family tree, as well as the excellent research being done by Lindsay Zanno and Peter Makovicky on the origin of plant-eating in theropod dinosaurs, and Falcarius starts to tell an interesting tail about the order in which therizinosaur traits appeared.

Falcarius appears to already be specialized for browsing for high forage. Given the lack of an enlarged gut for fermentation it probably preferred to seek out higher-quality plant matter, like fruiting bodies or seeds. The partially upright stance appears concurrently with a widening of the passage through the pelvis (not visible in side view) allowing move guts into that area, causing the center of gravity to sit further back despite the elongation of the neck.

The large hand claws (from which the authors derived the name "sickle-cutter") may have allowed Falcarius to pick up small prey, but they also may have served as defense for a fairly slow animal with small teeth. The first toe is low and long enough to start interacting with the ground, perhaps to provide balance and stability when browsing high.  All of these features would be carried to extremes in advanced therizinosaurs, but they seem to already be playing the same (albeit incipient) functional roles in Falcarius.

So with Falcarius we have an animal that at first glance appears inexplicably strange, but when viewed through the lens of where it was coming from (long-bodied small-headed meat eaters) and where it ends up (the upright, pot-bellied therizinosaurs) the combination of traits start to make a lot of sense.


Isn't science grand?


Kirkland, J. I., Zanno, L. E., Sampson, S. D., Clark, J. M. & DeBlieux, D. D., 2005. A primitive therizinosauroid dinosaur from the Early Cretaceous of Utah. Nature, v435, pp 84-87.

Zanno, L. E. 2006. The pectoral girdle and forelimb of the primitive therizinosauroid Falcarius utahensis (Theropoda, Maniraptora): Analyzing evolutionary trends withing Therizinosauroidea. Journal of Vertebrate Paleontology, v26 n3, pp 636-650.

Zanno, L. E. 2010. Osteology of Falcarius utahensis (Dinosauria: Theropoda): characterizing the anatomy of basal therizinosaurs. Zoological Journal of the Linnean Society. v158, pp 196-230.

Zanno, L. E. & Makovicky, P. J., 2011. Herbivorous ecomorphology and specialization patterns in theropod dinosaur evolution. Proceedings of the National Academy of Sciences. v108 m1, pp 232-237.

Um hey, Scientific American? Bird knees bend the same way as everyone else.


Ok, time for a quick anatomy lesson: Despite what you may have heard, bird knees do not bend backward.  Nor, in fact, do the knees of any tetrapod perform this trick.  Given the role of the knee in locomotion, it's not even clear how such a reversal could evolve after the initial "knee bend" direction was settled upon several hundred million years ago.

Why bring the anatomical equivalent of a fairy tale?  Well, it's a fairly common misconception.  So common, in fact, that it was recently enshrined by none other than Scientific American.  So let's see if we can clear this up with some simple diagrams:


Really, it's not.

It's a surprisingly common mistake.  When looking at living birds many people fail to realize that part of the leg is hidden on a bird; the upper leg barely moves, and along with the knee it is actually buried up under the feathers of the wing and body.  Birds also have quite long ankles, leaving their ankle joints in roughly the position we'd expect the knees to be on a human.  Like this:

The key here is that people are plantigrade animals, while all theropods (including birds) are digitigrade.  That means that human ankles are flat on the ground, and in our case our knees are roughly in the middle of our legs.  In birds and other digitigrade animals (most dinosaurs and many mammals, like dogs, deer, and horses) it's only the toes that contact the ground.  The ankle joint is well up off the ground, and the knee is is actually in the upper 1/3rd of the leg.  And in birds the thigh is actually even a smaller part of the leg, and as mentioned above is also mostly hidden under feathers.  Here's a look comparing the same leg portions of a human (in two poses), a dog, and an extinct bird:

(Presbyornis  copyright Scott Hartman, other skeletals modified after Charles Knight.)

(Presbyornis copyright Scott Hartman, other skeletals modified after Charles Knight.)

In the diagram the thigh bones (femora) are all colored red, the shin bones and proximal ankle bones are colored blue, the "foot bones" (the distal tarsals and the metatarsals) are green, and the individual toe bones are yellow.  Notice that the femur of the extinct bird Presbyornis is small and very high, and the knee (the joint between the red and blue bones) is up where it would be hidden by the body and wings.  But please also notice...the knee is bending the same way as ours.  And the same way as everything else that has a spine and walks on land.

Which brings us back to the article posted by Scientific American.  I don't want to go too far with "gotcha" blogging, but Scientific American is generally one of the more highly regarded popularizers of science on the web and in print.  I took a look the other pieces published by the author, and she seems like a solid reporter that just happens to have made a mistake.  Journalists don't get science degrees (and even if they did that would only be one subject), so this should not be construed as an attack.

But at the same time, this is a serious error.  It's like reporting that September has 32 days in it, or that the Red Sox clinched a playoff spot this week.  Only worse, as it's perpetuating a myth that gets passed around as "common knowledge".  I attempted to bring this to the attention of the relevant parties shortly after it was reported, both on Google+, where SciAm blog editor Bora Zivkovic has been making effective use of the new social network, and the author's Twitter account (which is frequently used).

Despite near real-time feedback, days have gone by with no correction. Making a mistake is understandable, but failing to correct it is not.  Scientific American has written hundreds of articles on the state of science education, and has often been an effective advocate for ways to improve it.  But the authority they derive comes from their attention to scientific detail, so I hope we will now see a quick correction without further delay.

Talos: A troodontid with a leg up on the competition


It's my pleasure to introduce the newest member of the troodontid family:Talos sampsoni. Named for paleontologist Scott Samspon, Talos was described by Lindsay Zanno and others in the wonderful open source journal Plos ONETalos is the first troodontid to be named from the Kaiparowits Formation of Utah, making it about 76 million years old.  Cross-sections of the long bones suggest that the animal was between four and six years old, and while it hadn't stopped growing, it appeared to be reaching reproductive age at a smaller size than it's close relative Troodon.

Of note is that the specimen had a bone in its second toe that was injured and partially healed.  Since it appears to have been injured from violent trauma, it's consistent with with the idea that the "switchblade" toe was used in a way that could result in such an injury (presumably either attack or defense).  Also, since the rest of the foot shows no indication of the sort of limping or other adjustments you see from a prolonged foot injury, it also reinforces the idea that troodontids walked with the second toe off the ground (as shown above).

I was producing the above skeletal reconstruction for the Utah Museum of Natural History, who has constructed a new building filled with exciting displays that will opened Fall 2012.  It should be a state of the art exhibit that everyone should go see if they get the chance.  Dr. Zanno was working on describing Talos at the time, so I worked with her to incorporate the data into the skeletal.  There is quite a bit of troodontid material from the Kaiparowits, but only a fraction of it can be confidently assigned toTalos at this time.  Restricting ourselves to the type specimen looks something like this:


 You could be forgiven if your first reaction is "that's not very complete!".  But I was thrilled.  Why?  It's hard to remember sometimes, since Troodon is illustrated frequently, but the published material that Troodon is usually based on is really scrappy.  Here is a quick and dirty example of what was published in Dale Russell's 1969 paper:


To create the reconstruction (then called Stenonychosaurus) ofTroodon

that inspired him to create his infamous dinosauroid sculpture, Dale Russell had to combine all of the material from the Asian taxa Saurornithoides (as seen below) just to create a usable composite:


That made the size of the head and pelvis more clear, and provided more back and tail vertebrae to scale from, but look at how "dotted" the outlines of the limbs are - there still wasn't a good guide for scaling the limbs of any of these advanced troodontids.  A few unpublished specimens are out there, but none of those provided definitive limb proportions. Talos finally does that.

So in addition to the interesting conclusions on biogeography, ontogeny, and mode of life, Talos should be exciting to illustrators because it provides the first definitive glimpse of the limb proportions of advanced troodontids.  They aren't radically different from the proportions Dale Russell speculated on in the 1960s (although the forelimbs are shorter than some people have speculated since then), but the good news is that we now know for sure.  In fact at this point the only serious unknown is the neck, although there are plenty of more basal troodonts to base that on.

Running around like an Ornitholestes with it's head cut off!


If you watched Episode II of Dinosaur Revolution, you may have laughed at this Ornitholetes, who I'll refer to as Ichabod.  This may seem like an odd scene to pick for a scientific discussion, but I think it actually has something useful to teach.  Also, I'm partially responsible.  I should be clear, the story idea was not mine (that's above my pay grade), but it's something that was run by me, and I did not try to shoot it down (and still wouldn't).

Why?  I think it's reasonable.

No wait, let me explain: I'll grant you that there isn't much in the professional literature on the subject on the subject of "headless running" in animals, but from some criticisms I've read I think people maybe thinking about this the wrong way (i.e., wondering about the distribution of "headless-running" in birds like the behavior it's an advanced condition).

Vertebrates as a group have one of the more centralized nervous systems among animals (with some arthropods and especially some cephalopods as the other contestants in the "flexibility over redundancy sweepstakes"), but tetrapod nervous system evolution in general is a story of progressive centralization that (so far) culminates in mammals.  Even in humans, with our gobs of ridiculously calorie-hungry centralized gray matter, we still have autonomous reactions that don't require the brain to be involved (as anyone knows who's burned themselves and jerked their hand away before they felt the pain).  That said, we have gone a long way down the path of nervous system centralization, and if you cut a mammal's head off you may get some twitching but it won't run around; our limbs literally cannot coordinate themselves without the brain's involvement (although morbidly it does appear that the head itself retains some coordination afterwards, if medieval reports are true that heads react for up to a 15 seconds after a beheading).

This degree of centralization is the derived condition, not the primitive one.  So it seems unlikely that chickens are special here, except in as much as they more frequently get clean beheadings in the presence of human observers than most other birds (a quick Google search shows that turkey's exhibit this as well).  This should be true of lizards, crocs, etc too (diapsids as a whole).  So with enough experimental trials I'd fully expect an Ornnitholestes to do a good headless chicken impression.  Now, I'll grant you that this would require a pretty clean bite on the allosaur's part, and the odds of observing it in the wild would not be very high.  But the sequence was devised as be a surprising bit of humor in a scenario that was possible, not probable.

Given those parameters it seems reasonable enough to me.


Back to our regularly scheduled blog posts...

Apologies for my multi-week absence.  I had to finish up some large projects.  I'm embarking some new cool ones as well, and I'll tell you about them as soon as I can.  In the mean time, I'll be getting back to posting blogs on dinosaur anatomy, as well as another series I have in mind on skeletal poses.  If you have any ideas of topics you'd like to see, please leave them below.  In the meantime, enjoy this skeletal drawing of Stokesosaurus, a tyrannosauroid that lived during the Late Jurassic of North America:

Tails of Woe


Welcome back!  This will be a shorter article that continues the concern for tails that we established in the inaugural T. rex tail post a couple weeks ago.  There is an all too common error that artists make when they attempt to impart a sense of to dynamic motion to their dinosaurs - and in particular to the dromaeosaurs.  They flex the tail up at the base so sharply that it would break the tail...if not break the pelvis!

Tail Tales: Break Dancing

We all like lively dancing dinosaurs; after a century of seeing moribund dinosaurs in swamps it's understandable that modern artists want to convey the "awesomeness" of their subjects.  If you ask me it can go too far sometimes - animals don't live their lives at 90 miles an hour - but we can all grasp the excitement of making a dynamic composition.  One way to impart motion is to have the tail doing something dramatic.  Alas, enhancing your dancing dinosaur this way without considering the anatomy may lead to an image where the animal has its tail disarticulated, or worse.

I mentioned dromaeosaur images tend to be among the worse offenders, and I know some of you are thinking "I've read that dromaeosaur tails can flex upwards at a 90 degree angle at the base of the tail!!!".  And it's true, the first several tail vertebrae are modified in such a way as to provide an expanded degree of flexibility (for up and down motion...not so much side to side), which means they have the ability to tilt the tail up sharply, to intimidate a rival, or just to better fit on your piece of paper.  But it's important to note that this tilt up takes place over the course of several vertebrae, meaning it can't happen like this:


At least it can't happen more than once unless the animal has good health insurance.  Now I certainly don't want to pick on Chris Srnka here - he's a fine artist and a lot of people make this mistake - this was just a great image to demonstrate the problem.  If you look at the image in handy-dandy X-ray format (as provided by Photoshop.

That red arrow is the steepest possible angle the tail could take emerging from the pelvis.  Why?  Dinosaur pelves have many vertebrae built into the sacrum (adding vertebrae to the sacrum is actually one of the characters that define what is and what isn't a dinosaur).  That sacrum fuses together and to the pelvis in adults, but even in juveniles there are no moving parts involved.


Front is to the left, the tail would be to the right (Carpenter & Wilson, 2008)

See?  There's just nothing that could move, even hypothetically.  In the case of dromaeosaurs the tail flexes up by spreading that 80 or so degrees of motion over 6+ joints, so none is flexing more than 14 degrees.  Here is a diagram of it:


So no more dromaeosaurs with tails growing out of their sacrum, please.

This isn't just a dromaeosaur problem either.  Many artists try to arch the tail base up on dinosaurs who don't naturally do this, and in so doing end up disarticulating the tail (or breaking the sacrum).  Even

Greg Paul's early Daspletosaurus painting fell prey to this temptation.

(Daspletosaurus, copyright Greg Paul, image from here.)

The problem is a bit more subtle in this painting, but the line of the vertebral column should extend gently down from the pelvis, while in this case it is flexed up right at the sacrum/tail juncture.  This would require a 25-30 degree flexure right at the first tail vertebrae (or else some flexing of the sacrum), which isn't going to happen without making the animal wince in sharp pain and reach for some Advil poste haste - and tyrannosaurs have horribly adapted arms for taking pain killers.

There are some important exceptions here.  Many sauropods, stegosaurs, and hadrosaurs have a bit of an upwards arch naturally at the tail base as it exits the pelvis.  Obviously those should have a bit of an arch (how much depends on the species in question).  But for most other dinosaurs, an arch of that magnitude isn't possible that immediately after the pelvis.


Sauropods like Mamenchisaurus have a natural flex in the tail base...but it still happens after the pelvis!

So remember, dinosaur tails may be flexible (depending on the group), but they aren't silly putty.  The vertebrae still need to articulate, and any motion you put into the tail needs to start after the hips, as the sacrum just can't bend.

Till next time, don't create your own tails of woe!


Barsbold, R. (1983). Carnivorous dinosaurs from the Cretaceous of Mongolia, Transactions of the Joint Soviet-Mongolian Paleontological Expedition v19, pp 5–119.

Carpenter, K. & Wilson, Y. (2008) A New Species of Camptosaurus (Ornithopoda: Dinosauria) from the Morrison Formation (Upper Jurassic) of Dinosaur National Monument, Utah, and a Biomechanical Analysis of Its Forelimb, Annals of Carnegie Museum, v76 n4, pp 227-263.