Giraffatitan's head is just WEIRD.

Posted by Nima On Monday, January 4, 2016 9 comments

The Jurassic weirdness continues!

The last post on Giraffatitan focused on the torso, and how it had often been inaccurately restored. But I didn't appreciate just how strange this animal's spine was until I got down to business, and started articulating images of the bones to see exactly how the centra and zygapohyses actually fit together. In the process I discovered that the actual 12th dorsal, though published by Werner Janensch in his 1950 monograph, was never scaled or reproduced in the mounted skeleton, nor was it used by any of the previous artists who had done skeletals of Giraffatitan. Greg Paul, Scott Hartman, Stephen Czerkas and or course (ironically) Janensch himself had left it out of their full-body skeletals. In addition it appears that they all changed the bizarre proportions of dorsal 9 - which has a relatively compact neural arch but a hugely elongated centrum - in order to make it fit in sequence such that the spine was more or less straight. But D9 (as heavily restored in plaster by Janensch anyway) has to be tilted upwards by around 40 degrees in order to have the short hyposphene reach far back enough to properly lock into place with D10, which indicates that their angle of articulation is anything but straight, and that D9 probably fits into the dorsal column like an upward-pointing wedge of sorts.

The result is a bizarre double-kink in the lower dorsals which both reinforces the lower back and makes the torso shorter and more compact. The fact that D10's centrum (again, going off of Janensch's restoration) has a condyle that is tilted up and back further reinforces this tilted angle of articulation demanded by the hyposphene of D9, as does the resulting snug fit of the neural spines of D9 and D10, without an excessive gap between them. Oddly Janensch doesn't carry over the weird shapes of both bones to his own full-body skeletal, though he does illustrate them individually in his paper, odd shapes and all, just it as his team restored them.

But this is far from the only strange thing about Giraffatitan that has been overlooked for decades.
Nearly every part of its body turned out to have unexpected features not included in ANY previous restorations. And one of the most commonly oversimplified, blurred, or just flat-out distorted parts in many restorations is... the head.

That's right, Giraffatitan's head is truly weird. A marvel of natural engineering and stress distribution through struts that in some places appear thinner than a human finger. The skull was light and hollow, yet could get up to a meter long (estimated size for adult individuals such as HMN XV2). And yet it was packed with big teeth resembling a cross between spoons and railroad spikes, built to crunch through hard branches high in the ancient conifers.

As you can see in the above picture, the skull is partially reconstructed with plaster, including one of the eye struts and the region just below the base of the nasal crest.

But there are in fact four skulls in existence. At least that is how many Janensch mentioned.

Three of them are missing a great deal of material, but the most well-known one, HMN t1, is nearly complete. We know this skull very well. Anyone who has seen photos of the Berlin mounted specimen (mostly based on HMN SII) has also probably seen this t1 skull, which is actually from a smaller individual. A scaled up cast of this skull was mounted on the skeleton itself in 2007, replacing an older crude sculpted skull.

The skull you see at the feet (or rather hands) of the Giraffatitan in these photos is actually only a cast of HMN t1. The real skull is stored in a museum vault and is (supposedly) off limits to the public.

Now at one point this face was cute.

Then it got fossilized and crushed. A few pieces such as the upper part of the eye socket are missing or broken. The upper jaw is partially collapsed in the middle, causing the sides of the maxillae to turn up and flare out. The sides of the jaws are thus artificially bowed out sideways. This led Dr. Matt Wedel to comment that it looks like a toilet seat today. Honestly I feel sorry for this poor creature. But sauropod skulls being delicate and easily smushed is a fact of life. Some of them had such loose connections between the skull bones that they actually dislocated during fossilization!

The crushing is easier to see from the side:


The snout has been flattened in the center, and to some extent the top of the nasal arch has also been squashed. Also notably, the teeth appear artificially long because they have slipped out of their sockets (or been pushed out by inclined crushing during fossilization) and the roots are visible. The skull itself had to be glued together from many fragments, and when first excavated was a bit of a jumble, like this:

Overall I would say given the shape the bones were in, Janensch and his staff had done a pretty good job of rebuilding the skull. Some of the skull elements were actually warped in the fossilization process, which makes sense as the bone layers are extremely thin.

But the trickiest aspect of this whole story is that there are a number of different ways the skull could have looked in real life. The crushing was uneven, which means the left and right sides of the skull appear rather different, with the right upper jaw considerably flatter than the left. Also we may be dealing with the possibility of ontogeny, that the skull of Giraffatitan would have changed shape with age and maturity. This is usually not a big concern in sauropods, as they do not develop any horns or massive butting surfaces on their heads, but that doesn't preclude the possibility that the shape of the head itself changed with age.

In trying to reconstruct a profile of Giraffatitan's head, I had to get around a few things.

First, the specimen I'm using for the skeletal is HMN SII, so the skull has to closely match the S116 skull, which is from the same or similar-sized individual. This skull has somewhat different proportions to some of the bones than HMN t1, although part of this may be due to either ontogeny or sex of the individual. That said, I wanted to create a reconstruction that adequately combines the most consistent aspects of all the skulls and eliminates crushing so that we can see the "ideal" morph of how SII's head would have looked on the living animal.

This was going to be a literal headache. It didn't help that Janensch and other early authors had themselves illustrated the "generic" Giraffatitan skull a number of different ways, with varying proportions.

So in brief, below, is the progress of morphs, trying to get the uncrushed proportions just exactly right (with a similar but shorter process for the referred Felch Quarry skull of Brachiosaurus - also an immature specimen - shown below it.)

By comparison with many photos from different angles and all the known Janensch engravings, gradually a more complete picture emerged. And so with a few remixes for different specimens, ultimately the conclusion was that the typical Giraffatitan head - hypothetically a mix of t1 and S116 - would look as follows.

So after about 30 variations and tweaks, this is what we've got. Overall a LOT better than the ugly derpy overbite version you see in most books and websites (basically a caricature of the crushed t1 skull), or for that matter the oversimplified blurry Greg Paul version which is sorely lacking in detail and deviates substantially from the fossils in several ways.

Pauly DERP that you can get sued for imitating. 

So yes, Giraffatitan - when uncrushed - has a rather different head than we've long been lead to believe. Feel free to comment below.

Dreadnoughtus - the Truth and the Myth

Posted by Nima On Saturday, September 13, 2014 4 comments

I knew this day was coming!

Lacovara's titanosaur has been described, named, and even 3D imaged! Dreadnoughtus schrani, read about it here: .

I saw this animal in preparation at the lab in Carnegie Museum long before it had a name, and I had the opportunity to speak with Dr. Lacovara himself at SVP 2010 and 2011 about this animal's morphology and likely lineage, so seeing the fruits of the Drexel team's work is personally satisfying. And unlike many instances of new dinosaur names, this one fits the bill perfectly. Dreadnoughtus is indeed a dreadnought of the dinosaurs. Not the only one by any means, but it's about time sauropods got some really imposing-sounding names, not just predatory theropods like T. rex. At some point sauropods really do get too big for any predator to mess with.

Now as usual with these giant dinosaur discoveries there are a number of myths floating around with the facts. Some of the most common rumors media reports that we can scrutinize are as follows:

1. This is the biggest dinosaur yet found.
Actually, no it's not. Argentinosaurus, Alamosaurus, Puertasaurus, Ruyangosaurus, the Chubut Monster, "Huanghetitan" ruyangensis, and maybe even Paralititan and Argyrosaurus sp. likely outclass it for raw size and mass. And there are a few brachiosaurs which also might be larger, including a very big referred specimen of Brachiosaurus itself. Of course neither specimen of Dreadnoughtus is fully grown (unfused scapula-coracoid suture is a dead giveaway of immaturity) so perhaps an adult Dreadnoughtus could rank higher in the "top 10 biggest". It is however among the biggest dinosaurs that are actually complete enough to reasonably estimate their size without much room for guesswork. But nobody is officially claiming it to be the "biggest" - we know based on general large titanosauriform proportions that most or all of the aforementioned animals are larger, even if estimating just how much larger is a bit more tricky.

2. This is the most complete giant titanosaur found.
This is technically true. Although it's missing most of the neck and ribs, which are largely present in Futalognkosaurus, the other contender for "most complete giant titanosaur". The fully tally of bones from the two Dreadnoughtus specimens adds up to more, especially considering the limb material, but the published diagrams and 3D scans still look less complete than Futalognkosaurus - when you composite all 3 specimens of Futa together, anyway. Surface area-wise, Futa's huge and deep neck bones at least make it appear more complete. And the referred juveniles contribute some limb material. Both animals appear to be similar in size.

3. This animal is a lognkosaur, as the paper seems to imply.
What's funny is that when the fossils were still under preparation back in 2010-11, both Dr. Lacovara and myself thought it might be something much more derived, like a record-sized Aeolosaurus-grade saltasaurid, due to the forward-slanted, antenna-like prezygapophyses in the tail. Of course to be fair, most of the fossils were in fragments at the time and had to be painstakingly glued back together, and when part or all of a caudal neural spine is broken off, many unrelated titanosaurs can look deceptively like "aeolosaurs". However, after analyzing the paper I can say that a lognkosaur-like position is indeed more plausible. This animal however isn't quite a lognkosaur. Close, but not quite.
The upper humerus bulges upward medially, much more like Argyrosaurus and Quetecsaurus, which has a very Argyrosaurus-like hand shape. Unfortunately hand material is missing for Dreadnoughtus, but the arm bones look Argyrosaur-like enough. The femur is very wide and robust with a large, midlevel 4th trochanter, convex-curved upper margin, and a reduced lateral bulge - very different from any known lognkosaur femur, but very similar to the two huge "Argyrosaurus sp." femurs in the Field Museum. The dorsal vertebrae are pretty wide with substantial laminae making the neural spines triangular, but just not as wide as in true lognkosaurians. They do however bear a passing resemblance to the dorsals of "Argyrosaurus sp." (=Elaltitan lilloi) specimen PVL 4628. The paper clusters it with Malawisaurus, which it places in a more derived position than true lognkosaurs - a position that isn't supported by the humerus and femur morphology and think should be re-examined. The Paleo King ranks this species as an Argyrosaurid, probably closest to Quetecsaurus. Which is a very good thing indeed, as it (along with the even odder Quetecsaurus) greatly improves our knowledge of this obscure titanosaur family.

Bones NOT to scale here!

Or here, though looks may deceive!

4. The neck was horizontal (?!?!?!?)
The reality is that only two of the neck bones were found, and both exhibit pretty serious crushing. And the first 3 dorsal vertebrae are missing entirely, with the 4th being seriously crushed as well. So it's very difficult if not impossible to reconstruct its neck posture accurately. The honest answer is that we don't know what Dreadnoughtus' neck posture was. But I think it's a pretty good guess that the neck was vertical or semi-vertical as in Futalognkosaurus, not horizontal as in the published skeletal in the paper. We are definitely dealing with a high-browser here. Titanosaurs of such large size usually have the extremely long and vertical necks of high browsers - there is no point in wasting such necks to only bend down and eat ferns like Saltasaurus or Diplodocus. Even the paltry tooth material found with Dreadnoughtus is that of a high-browsing conifer eater. The 9th(?) cervical is big enough to indicate an even longer neck than Futalognkosaurus, and even with the 3D model omitting any speculative reconstruction of the hands (and leaving precious little space for them), the shoulders of Dreadnoughtus are still higher than its hips. Add in hands of correct lognkosaur or argyrosaur proportions, and this will tilt up the torso and shoulders even further. And the general rule with all high-shouldered sauropods is that they also tend to be high-browsers, regardless of neck length. So we have two patterns of evidence in favor of a vertical neck.

3d model scan of the actual fossils. Even with the arm not raised high enough to leave space for the hands, the shoulders rise above the hips. Every well-preserved macronarian with this pattern exhibits a more or less vertical neck.  Note that the back is pretty straight here.
Original skeletal from paper - the neck seems to be horizontal to make room for more figures? The back is oddly curved, which isn't evidence by anything in the fossils, and the dorsal spines just look wrong. Where's the backsweep?
Very hastily corrected version with correct-sized hands and a straighter back and more vertical neck. Now that's better!
Unfortunately these days (thanks in no small part to a certain wispy white mustache from outside the field) the convention in many dinosaur papers seems to be to illustrate sauropod necks horizontal since it's seen as more "conservative" and "safe" even if it isn't meant to be literally accurate or isn't the main focus of the paper. Never mind that it wastes neck length, puts much more strain on the span of the neck than a vertical posture, and that most macronarian necks weren't built for tolerating horizontal strain for any substantial length of time... of course it's possible Lacovara and co. may have just wanted to save vertical space per page in an already large paper (publishing is getting expensive as many of us know) so I can't judge their reasoning on this particular case.

5. Dreadnoughtus and Puertasaurus were basically best chums.
Doubtful. Although they were found very close together (something like only 13 miles apart) in Santa Cruz province in the far south of Argentina, their time horizons may be different. The exact stratigraphy of the Dreadnoughtus site is still a bit murky, and even if it is Maastrichtian in age, it may still be separated from Puertasaurus by hundreds of thousands of years if not more. And even if they did live at the same time, they may not have been buddies. Big sauropods in similar feeding niches were just as likely to compete over resources as share them. Notice that term similar niches. This did not mean that big herbivores always butted heads (or tails, as the case may be). Sauropods in vastly different feeding niches (such as brachiosaurs and diplodocids in the Morrison and Tendaguru formations) tended to coexist just fine.

Plus, if the subadult type specimen of Dreadnoughtus (or one of similar size, around 50-60 tons) crossed paths with the Puertasaurus type (around 90-110 tons, which may or may not be an adult) then maybe "dread-nought" may have stopped being such an appropriate label! Comparing the dorsals of Dreadnoughtus with the one found from Puertasaurus, you will soon see that while Dreadnoughtus may be huge, it's definitely not in the same league.

Is the French Monster a Euhelopodid?

Posted by Nima On Thursday, July 10, 2014 0 comments

One of these days we may see a paper come out about this. France, as you may know, contains a dinosaur of truly gigantic proportions (and no, I'm not talking about the Algerian species "Brachiosaurus" nougaredi which seems to have disappeared into a black hole somewhere in Paris). No, this one's a local. A new "titanosaur" known only as the French Monster, or the giant of Angeac, has turned up in the past few years in a bone bed which contains a huge jumble of Early Cretaceous dinosaurs of various sizes - everything from baby abelisaurs to massive hundred-foot sauropods.

Except it's not really a titanosaur, is it?

The French Monster is another one of those legendary or semi-legendary sauropods which should have a name and a formal description, though it's unlikely most of them ever will. Even the pictures evoke something rarely seen in the fossil record.

Mainly the image that sticks in your mind is the very long and oddly blackened right femur, over 2.2m long, with which everyone seems to be "doing the Jensen" though none of them come close to getting it right.

But there is much more to the Angeac-Charente site where this bone was found. There appears to be part of a second femur also recovered at the site, and possible rib fragments. And different individuals of this species have been found, with the fossils in different shades of mineralization.

GIGANTIC tail vertebrae and toe bones. Over a foot in length or diameter. Note the cheesy scaled-up mid-90s "Brachiosaurus" skeleton model designed by Dale Russell and Ely Kish - these things originally came in a screw-cap plastic egg, and were the highlight of every kid's otherwise dull-as-doorknobs trip to T.J. Maxx. Some of you may be too young to remember... The model isn't 6 feet tall like it may appear, it's on the shelf, not the floor!

The heavy toe bones and caudals. These may be from multiple individuals.

Some teeth were found, very well preserved and encrusted with mineral deposits formed in fossilization.

These teeth have a very basal appearance, and so the labeling of this animal as a titanosaur is unlikely. The teeth could easily pass for Brachiosaur teeth. But there is more than just this first indication what what we are seeing may not be a true titanosaur.

The femur and the other pieces were apparently packed and shipped to a museum. It's not clear if this is in Paris or elsewhere. The femur is crushed and snapped in a few places but its total length is still preserved. A cast and wall mount were rapidly made. And this is the peculiar part.

Here you see the femur, fragments of another, and a rib. Apparently this is a part of the full wall mount you see in the background of the previous picture. It's evident both that there was crushing and that very little erosion has happened. The most unusual things about this femur are the very long femoral head, and the odd lower end, with its inner condyle extending lower than the outer one. This is the exact opposite of when you see in derived titanosaurs. But is is a classic femur morph in a group which is close to basal titanosaurs.

Between brachiosaurs and basal somphospondyli (creatures like Chubutisaurus, Ligabuesaurus, and some would say, Paluxysaurus and Sauroposeidon) and true titanosaurs, there was an amazing radiation of transitional forms. These appear to form at least two major families: Euhelopodidae, and Acrofornica (tall-arches). Both are characterized by extremely long necks, high cervical counts, and bifid neural spines in the neck. The Acrofornica are further distinguished by very tall neural arches, high diapophyses, and nearly no neural spine in the dorsals. They tend to have well-separated sacral ribs, whereas those of euhelopodids tend to be extensively fused together. And whereas euhelopodids (or at least some of them) have procoelous tail vertebrae (anticipating derived titanosaurs!) the tails of acrofornicans revert back to simple amphiplatyan tails, as in basal titanosaurs.

After about 3 years of morphometric comparisons between various elements and overlaps between different specimens (some of them very fragmentary) the following family tree slowly began to reveal itself. This is not a complete family tree of titanosauriformes; only some of the more well-known ones intermediate between brachiosaurs and titanosaurs are included here.

And in the comparisons between different elements, specifically femora, the French Monster appears to belong in this transitional "gap" rather than in the more derived true Titanosauria. Previously I had put several hypotheses out there, what the French Monster may have been. Perhaps a lognkosaur, possibly a Malawisaurus-grade stem lognkosaur. Perhaps an oddly slender Antarctosaur of Pellegrinisaurus ilk. Maybe even a nemegtosaurid like Huabeisa- wait a minute, that one's not even considered a nemegtosaur anymore! That's the great thing about finally using digital photos... they clear up all sorts of confusion - which Pang & Cheng had forgotten to do in the original blurry description.

Well it turns out that the French Monster is more like Huabeisaurus than previously realized. Both are closer to Euhelopus and the acrofornicans than to true titanosaurs.

Complicating matters is the fact that some lognkosaurian titanosaurs have a protruding femoral head and a high and prominent lateral bulge, which differs from all other titanosaurs, and converges on that of some euhelopodids and acrofornicans. However the distal end of the femur follows radically different patterns in the two lineages. Here's the comparison of posterior views, you be the judge (not to scale):

So far the morphology of the French Monster's femur (the long and slender femoral head, the tibial condyle being longer and more massive than the fibular condyle, the lateral bulge high on the femur) appear more like a basal somphospondyl than a titanosaur. Most specifically, it resembles Paluxysaurus, and to a lesser degree, Huabeisaurus. The distal end of the Chubutisaurus femur is too eroded to make a good comparison on the condyles. All the same, the basal lineage of this unusual animal look much clearer now. The French Monster points to a radiation of basal sonphospondyli well outside Asia in the early Cretaceous. Angolatitan was the first euhelopodid to be described outside Asia, proving that the clade was not an exclusively Chinese one - and there are rumors of some similar fossils in Europe as well. If it's a chubutisaur, it would be the first to be found outside the Americas.

Perhaps the biggest oddity of all is how slender the French Monster's femur is. The crushing is mostly from front to back, not lateral. So it really was this narrow. There is a possibility then, that this animal was not even close to the maximum size possible. As an adult it may have been more robust. There is no coracoid or scapula material, so the degree of suture fusion in the shoulders (and thus the animal's maturity) is open to speculation.

As far as I can see it, this dinosaur is most likely a chubutisaur (or whatever Paluxysaurus is, seeing as it's more or lesss totally busted as a purported brachiosaur), but may also belong in acrofornica or euhelopodidae depending on how the cladistics stack up. In any case it's huge and unusual, and maybe soon we may get to see a description and some idea of its overall proportions.