February 15, 2014
“Look at all the things you’ve done for me
Opened up my eyes,
Taught me how to see,
Notice every tree.”
So sings Dot in Move On, the climactic number of Stephen Sondheim’s Pulitzer Prize-winning music Sunday in the Park with George, which on the surface is about the post-impressionist painter Georges Seurat, but turns out to be a study of obsession and creativity.“Taught me how to see”? What kind of talk is that? One the surface, it seems silly — we all know how to see. We do it constantly, without thinking. Yet it’s something that artists talk about all the time. And anyone who’s sat down and seriously tried to paint or draw something will have some understanding of what the phrase means. We have such strong implicit ideas of what things look like that we tend to reproduce what we “know” is there rather than what’s actually there. Like I said, we see without thinking.
In fact, the psychology of perception is complicated and sophisticated, and the brain does an extraordinary amount of filtering of the visual signals we get, to save us the bother of having to consciously process way too much data. This is a whole scientific field of its own, and I’m going to avoid saying very much about it for fear of making a fool of myself — as scientists so often do when wandering outside their own field. But I think it’s fair to say that we all have a tendency to see what we expect to see.
In the case of sauropods, this tendency has meant that we’ve all been startlingly bad at seeing pneumaticity in the caudal vertebrae of sauropods. Because the literature has trained us to assume it’s not there. For example, in the two competing sauropod phylogenies that dominated the 2000s, both Wilson (2002) and Upchurch et al. (2004) scored caudal pneumaticity as very rare: Wilson’s character 119, “Anterior caudal centra, pneumatopores (pleurocoels)”, was scored 1 only for Diplodocus and Barosaurus; and Upchurch et al. (2004:286) wrote that “A few taxa (Barosaurus, Diplodocus, and Neuquensaurus) have pleurocoel-like openings in the lateral surfaces of the cranial [caudal] centra that lead into complex internal chambers”. That’s all.
And that’s part of the reason that every year since World War II, a million people have walked right past the awesome mounted brachiosaur in the Museum Für Naturkunde Berlin without noticing that it has pneumatic caudals. After all, we all knew that brachiosaur caudals were apneumatic.
But in my 2005 Progressive Palaeontology talk about upper limits on the mass of land animals estimated through the articular area of limb-bone cartilage, I included this slide that shows how much bigger the acetabulum of Giraffatitan is than the femoral head that it houses:
And looking at that picture made me wonder: those dark areas on the sides of the first few caudals (other than the first, which is a very obvious plaster model) certainly look pneumatic.
Then a few years later, I was invited to give a talk at the Museum Für Naturkunde Berlin itself, on the subject “Brachiosaurus brancai is not Brachiosaurus“. (This of course was drawn from the work that became my subsequent paper on that subject, Taylor 2009) And as I was going through my photos to prepare the slides of that talk, I thought to myself: darn it, yes, it does have pneumatic caudals!
So I threw this slide into the talk, just in passing:
Those photos were pretty persuasive; and a closer examination of the specimen on that same trip was to prove conclusive.
Earlier in 2009, I’d been in Providence, Rhode Island, with my Index Data colleagues. I’d managed to carve a day out of the schedule to hope along the coast to the Yale Peabody Museum in New Haven, Connecticut. My main goal was to examine the cervicals of the mounted Apatosaurus (= “Brontosaurus“) excelsus holotype (although it was also on that same trip that I first saw the Barosaurus holotype material that we’ve subsequently published a preprint on).
The Brontosaurus cervicals turned out to be useless, being completely encased in plaster “improvements” so that you can’t tell what’s real and what’s not. hopefully one day they’ll get the funding they want to take that baby down off its scaffold and re-prep the material.
But since I had the privilege of spending quality time with such an iconic specimen, it would have been churlish not to look at the rest of it. And lo and behold, what did I see when I looked at the tail but more pneumaticity that we thought we knew wasn’t there!
What does this mean? Do other Giraffatitan and Apatosaurus specimens have pneumatic tails? How pervasive is the pneumaticity? What are the palaeobiological implications?
Stay tuned! All will be revealed in Matt’s next post (or, if you can’t wait, in our recent PLOS ONE paper, Wedel and Taylor 2013b)!
- Taylor, Michael P. 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914). Journal of Vertebrate Paleontology 29(3):787-806.
- Upchurch, Paul, Paul M. Barrett and Peter Dodson. 2004. Sauropoda. pp. 259-322 in D. B. Weishampel, P. Dodson and H. Osmólska (eds.), The Dinosauria, 2nd edition. University of California Press, Berkeley and Los Angeles. 861 pp.
- Wedel, Mathew J., and Michael P. Taylor. 2013. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus. PLOS ONE 8(10):e78213. 14 pages. doi:10.1371/journal.pone.0078213
- Wilson, J.A. 2002. Sauropod dinosaur phylogeny: critique and cladistic analysis. Zoological Journal of the Linnean Society 136:217-276.
January 15, 2014
[This is part 4 in an ongoing series on our recent PLOS ONE paper on sauropod neck cartilage. See also part 1, part 2, and part 3.]
Here’s a frequently-reproduced quote from Darwin:
About thirty years ago there was much talk that geologists ought only to observe and not theorise; and I well remember some one saying that at this rate a man might as well go into a gravel-pit and count the pebbles and describe the colours. How odd it is that anyone should not see that all observation must be for or against some view if it is to be of any service!
It’s from a letter to Henry Fawcett, dated September 18, 1861, and you can read the whole thing here.
I’ve known this quote for ages, having been introduced to it at Berkeley–a copy used to be taped to the door of the Padian Lab, and may still be. It’s come back to haunt me recently, though. An even stronger version would run something like, “If you don’t know what you’re looking for, you won’t make the observation in the first place!”
For example: I started CT scanning sauropod vertebrae with Rich Cifelli and Kent Sanders back in January, 1998. Back then, I was interested in pneumaticity, so that’s what I looked for, and that’s what I found–work which culminated in Wedel et al. (2000) and Wedel (2003). It wasn’t until earlier this year that I wondered if it would be possible to determine the spacing of articulated vertebrae from CT scans. So everything I’m going to show you, I technically saw 15 years ago, but only in the sense of “it crossed my visual field.” None of it registered at the time, because I wasn’t looking for it.
A corollary I can’t help noting in passing: one of the under-appreciated benefits of expanding your knowledge base is that it allows you to actually make more observations. Many aspects of nature only appear noteworthy once you have a framework in which to see them.
So anyway, the very first specimen we scanned way back when was the most anterior of the three plaster jackets that contain the four cervical vertebrae that make up OMNH 53062, which was destined to become the holotype of Sauroposeidon. I’ve written about the taphonomy of that specimen here, and you can read more about how it was excavated in Wedel and Cifelli (2005). We scanned that jacket first because, although the partial vertebrae it contains are by far the most incomplete of the four, the jacket is a lot smaller and lighter than the other two (which weigh hundreds of pounds apiece). Right away we saw internal chambers in the vertebrae, and that led to all of the pneumaticity work mentioned above.
Happily for me, that first jacket contains not only the posterior two-thirds of the first vertebra (possibly C5), but also the front end of the second vertebra. Whoever decided to plow through the second vertebra to divide the specimen into manageable chunks in the field made a savvy choice. Way back in 2004 I realized that the cut edge of the second vertebra was not obscured by plaster, and therefore the internal structure could be seen and measured directly, which is a lot cleaner than relying on the artifact-heavy CT scans. (The CT scans are noisy because the hospital machines we had access to start to pant a bit when asked to punch x-rays through specimens this large and dense.) A figure derived from that work made it into a couple of papers and this post, and appears again above.
But that’s pneumaticity, which this post is allegedly not about. The cut through the second vertebra was also smart because it left the intervertebral joint intact.
Here are a photo of the jacket and a lateral scout x-ray. The weird rectangles toward the left and right ends of the x-ray are boards built into the bottom of the jacket to strengthen it.
And here’s a closeup of the C5/C6 joint, with the relevant radiographs and tracing. The exciting thing here is that the condyle is centered almost perfectly in the cotyle, and the zygapophyses are in articulation. Together with the lack of disarticulation in the cervical rib bundle (read more about that here and in Wedel et al. 2000), these things suggest to us that the vertebrae are spaced pretty much as they were in life. If so, then the spacing between the vertebrae now tells us the thickness of the soft tissue that separated the vertebrae in life.
I should point out here that we can’t prove that the spacing between the vertebrae is still the same as it was in life. But if some mysterious force moved them closer together or farther apart, it did so (1) without decentering the condyle of C6 within the cotyle of C5, (2) without moving the one surviving zygapophyseal joint out of contact, and (3) without disarticulating the cervical ribs. The cervical ribs were each over 3 meters long in life and they formed vertically-stacked bundles on either side below the vertebrae; that’s a lot of stuff to move just through any hypothetical contraction or expansion of the intervertebral soft tissues after death. In fact, I would not be surprised if the intervertebral soft tissues did contract or expand after death–but I don’t think they moved the vertebrae, which are comparatively immense. The cartilage probably pulled away from the bone as it rotted, allowing sediment in. Certainly every nook and cranny of the specimen is packed with fine-grained sandstone now.
Anyway, barring actual preserved cartilage, this is a best-case scenario for trying to infer intervertebral spacing in a fossil. If articulation of the centra, zygs, and cervical ribs doesn’t indicate legitimate geometry, nothing ever will. So if we’re going to use the fossils to help settle this at all, we’re never going to have a better place to start.
So, by now, you know I’m a doofus. I have been thinking about this problem literally for years and the data I needed to address it was sitting on my hard drive the entire time. One of the things I pondered during those lost years is what the best shape for a concave-to-convex intervertebral joint might be. Would the best spacing be radially constant (A in the figure above), or antero-posteriorly constant (B), or some other, more complicated arrangement? The answer in this case surprised me–although the condyle is a lot smaller in diameter than the cotyle, the anteroposterior separation between them in almost constant, as you can see in part C of the above figure.
Don’t get too worked up about that, though, because the next joint is very different! Here’s the C6/C7 joint, again in a lateral scout x-ray, with the ends of the bones highlighted. Here the condyle is almost as big in diameter as the cotyle, but it is weirdly flat. This isn’t a result of overzealous prep–most of the condyle is still covered in matrix, and I only found its actual extent by looking at the x-ray. This is flatter than most anterior dorsal vertebrae of Apatosaurus–I’ve never seen a sauropod cervical with such a flat condyle. Has anyone else?
The condyle of C6 is a bit flatter than expected, too–certainly a lot flatter than the cervical condyles in Giraffatitan and the BYU Brachiosaurus vertebrae. As we said in the paper,
It is tempting to speculate that the flattened condyles and nearly constant thickness of the intervertebral cartilage are adaptations to bearing weight, which must have been an important consideration in a cervical series more than 11 meters long, no matter how lightly built.
Anyway, obviously here the anteroposterior distance between condyle and cotyle could not have been uniform because they are such different shapes. Wacky. The zygs are missing, so they’re no help, and clearly the condyle is not centered in the cotyle. Whether this posture was attainable in life is debatable; I’ve seen some pretty weird stuff. In any case, we didn’t use this joint for estimating cartilage thickness because we had no reason to trust the results.
Kent Sanders and I had also scanned several of the smaller sauropod vertebrae from the Carnegie collection (basically, the ones that would fit in the trunk of my car for the drive back to Oklahoma). Crucially, we’d scanned a couple of sets of articulated vertebrae, CM 3390 and CM 11339, both from juvenile individuals of Apatosaurus. In both cases, the condyles and cotyles are concentric (that’s what the ‘orthogonal gaps’ are all about in the above figure) and the zygs are in articulation, just as in Sauroposeidon. These are dorsals, so we don’t have any cervical ribs here to provide a third line of evidence that the articulation is legit, but all of the evidence that we do have is at least consistent with that interpretation.
So, here’s an interesting thing: in CM 3390, above, the first dorsal is cranked up pretty sharply compared to the next one, but the condyle is still centered in the cotyle and the zygs are in articulation. Now, the vertebrae have obviously been sheared by taphonomic deformation, but that seems to have affected both vertebrae to the same extent, and it’s hard to imagine some kind of taphonomic pressure moving one vertebra around relative to the next. So I think it’s at least plausible that this range of motion was achievable in life. Using various views and landmarks, we estimate the degree of extension here somewhere between 31 and 36 degrees. That’s a lot more than the ~6 degrees estimated by Stevens and Parrish (1999, 2005). And, as we mentioned in the paper, it nicely reinforces the point made by Upchurch (2000), that flexibility in the anterior dorsals should be taken into account in estimating neck posture and ROM.
Here’s our last specimen, CM 11339. No big surprises here, although if you ever had a hard time visualizing how hyposphenes and hypantra fit together, you can see them in articulation in parts C and D (near the top of the specimen). Once again, by paging through slices we were able to estimate the separation between the vertebrae. Incidentally, the condyle IS centered in the cotyle here, it just doesn’t look that way because the CT slice is at an angle to the joint–see the lateral scout in part A of the figure to see what I mean.
So, what did we find? In Sauroposeidon the spacing between C5 and C6 is 52mm. That’s pretty darn thick in absolute terms–a shade over two inches–but really thin in relative terms–only a little over 4% of the length of each vertebra. In both of the juvenile Apatosaurus specimens, the spacing between the vertebrae was about 14mm (give or take a few because of the inherent thickness of the slices; see the paper for details on these uncertainties).
Now, here’s an interesting thing: we can try to estimate the intervertebral spacing in an adult Apatosaurus in two ways–by scaling up from the juvenile apatosaurus, or by scaling sideways from Sauroposeidon (since a big Apatosaurus was in the same ballpark, size-wise)–and we get similar answers either way.
Scaling sideways from Sauroposeidon (I’m too lazy to write anymore so I’m just copying and pasting from the paper):
Centrum shape is conventionally quantified by Elongation Index (EI), which is defined as the total centrum length divided by the dorsoventral height of the posterior articular surface. Sauroposeidon has proportionally very long vertebrae: the EI of C6 is 6.1. If instead it were 3, as in the mid-cervicals of Apatosaurus, the centrum length would be 600 mm. That 600 mm minus 67 mm for the cotyle would give a functional length of 533 mm, not 1153, and 52 mm of cartilage would account for 9.8% of the length of that segment.
Scaling up from the juveniles: juvenile sauropods have proportionally short cervicals (Wedel et al. 2000). The scanned vertebrae are anterior dorsals with an EI of about 1.5. Mid-cervical vertebrae of this specimen would have EIs about 2, so the same thickness of cartilage would give 12mm of cartilage and 80mm of bone per segment, or 15% cartilage per segment. Over ontogeny the mid-cervicals telescoped to achieve EIs of 2.3–3.3. Assuming the cartilage did not also telescope in length (i.e., didn’t get any thicker than it got taller or wider), the ratio of cartilage to bone would be 12:120 (120 from 80*1.5), so the cartilage would account for 10% of the length of the segment–almost exactly what we got from the based-on-Sauroposeidon estimate. So either we got lucky here with our tiny sample size and truckloads of assumptions, or–just maybe–we discovered a Thing. At least we can say that the intervertebral spacing in the Apatosaurus and Sauroposeidon vertebrae is about the same, once the effects of scaling and EI are removed.
Finally, we’re aware that our sample size here is tiny and heavily skewed toward juveniles. That’s because we were just collecting targets of opportunity. Finding sauropod vertebrae that will fit through a medical-grade CT scanner is not easy, and it’s just pure dumb luck that Kent Sanders and I had gotten scans of even this many articulated vertebrae way back when, since at the time we were on the hunt for pneumaticity, not intervertebral joints or their soft tissues. As Mike has said before, we don’t think of this paper as the last word on anything. It is, explicitly, exploratory. Hopefully in a few years we’ll be buried in new data on in-vivo intervertebral spacing in both extant and extinct animals. If and when that avalanche comes, we’ll just be happy to have tossed a snowball.
- Stevens, K.A. and Parrish, J.M. 1999. Neck posture and feeding habits of two Jurassic sauropod dinosaurs. Science 284: 798-800. [Free subscription required]
- Stevens, Kent A., and J. Michael Parrish. 2005. Neck posture, dentition, and feeding strategies in Jurassic sauropod dinosaurs. pp. 212-232 in: Virginia Tidwell and Ken Carpenter (eds.), Thunder Lizards: the Sauropodomorph Dinosaurs. Indiana University Press, Bloomington, Indiana. 495 pp.
- Taylor, Michael P., and Mathew J. Wedel. 2013c. The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. PLOS ONE 8(10): e78214. 17 pages. doi:10.1371/journal.pone.0078214 [PDF]
- Upchurch, P. 2000. Neck posture of sauropod dinosaurs. Science 287: 547b.
- Wedel, M.J. 2003b. The evolution of vertebral pneumaticity in sauropod dinosaurs. Journal of Vertebrate Paleontology 23:344-357.
- Wedel, M.J. 2007. Aligerando a los gigantes (Lightening the giants). ¡Fundamental! 12:1-84. [in Spanish, with English translation]
- Wedel, M.J., and Cifelli, R.L. 2005. Sauroposeidon: Oklahoma’s native giant. Oklahoma Geology Notes 65 (2):40-57.
- Wedel, M.J., R.L. Cifelli and R.K. Sanders. 2000. Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica 45(4): 343-388.
October 31, 2013
First, there’s a nice write-up of one of our papers (Wedel and Taylor 2013b on pneumaticity in sauropod tails) in the Huffington Post today. It’s the work of PLOS blogger Brad Balukjian, a former student of Matt’s from Berkeley days. The introduction added by the PLOS blogs manager is one of those where you keep wanting to interrupt, “Well, actually it’s not quite like that …” but the post itself, once it kicks in, is good. Go read it.
Brad also has a guest-post on Discover magazine’s Crux blog: How Brachiosaurus (and Brethren) Became So Gigantic. He gives an overview of the sauropod gigantism collection as a whole. Well worth a read to get your bearings on the issue of sauropod gigantism in general, and the new collection in particular.
PLOS’s own community blog EveryONE also has its own brief introduction to the collection.
And PLOS and PeerJ editor Andy Farke, recently in these pages because of his sensational juvenile Parasaurolophus paper, contributes his own overview of the collection, How Big? How Tall? And…How Did It Happen?
Finally, if you’re at SVP, go and pick up your free copy of the collection. Matt was somehow under the impression that the PLOS USB drives with the sauropod gigantism collection would be distributed with the conference packet when people registered. In fact, people have to go by the PLOS table in the exhibitor area (booth 4 in the San Diego ballroom) to pick them up. There are plenty of them, but apparently a lot of people don’t know that they can get them.
- Wedel, Mathew J., and Michael P. Taylor. 2013. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus. PLOS ONE 8(10):e78213. 14 pages. doi:10.1371/journal.pone.0078213 [PDF]
October 30, 2013
This is an exciting day: the new PLOS Collection on sauropod gigantism is published to coincide with the start of this year’s SVP meeting! Like all PLOS papers, the contents are free to the world: free to read and to re-use. (What is a Collection? It’s like an edited volume, but free online instead of printed on paper.)
There are fourteen papers in the new Collection, encompassing neck posture (yay!), nutrition (finally putting to bed the Nourishing Vomit Of Eucamerotus hypothesis), locomotion, physiology and evolutionary ecology. Lots every sauropod-lover to enjoy.
Matt and I are particularly excited that we have two papers in this collection: Taylor and Wedel (2013c) on intervertebral cartilage in necks, and Wedel and Taylor (2013b) on pneumaticity in the tails of (particularly) Giraffatitan and Apatosaurus. So we have both ends of the animal covered. It also represents a long-overdue notch on our bed-post: for all our pro-PLOS rhetoric, this is the first time either of has had a paper published in a PLOS journal.
It’s a bit of a statistical anomaly that after a decade of collaboration in which there was never a Taylor & Wedel or Wedel & Taylor paper, suddenly we have five of them out in a single year (including the Barosaurus preprint, which we expect to eventually wind up as Taylor and Wedel 2014). Sorry about the alphabet soup.
Since Matt is away at SVP this week, I’ll be blogging mostly about the Taylor and Wedel paper this week. When Matt returns to civilian life, the stage should be clear for him to blog about pneumatic caudals.
- Taylor, Michael P., and Matthew J. Wedel. 2013c. The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. PLOS ONE 8(10): e78214. 17 pages. doi:10.1371/journal.pone.0078214 [PDF]
- Wedel, Mathew J., and Michael P. Taylor. 2013. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus. PLOS ONE 8(10):e78213. 14 pages. doi:10.1371/journal.pone.0078213 [PDF]
September 11, 2013
We’ve blogged a lot of Bob Nicholls‘ art (here, here, and here) and we’ll probably continue to do so for the foreseeable future. We don’t have much choice: he keeps drawing awesome things and giving us permission to post them. Like this defiantly shaggy Apatosaurus, which was probably the star of the Morrison version of Duck Dynasty. Writes Bob:
On my way home at the airport I did a sketch of your giant Apatosaurus* — see attachment. My thought was that massive thick necks were probably pretty sexy things to apatosaurs, so maybe sexually mature individuals used simple feathers (stage 1, 2 or 3?) to accentuate the neck profile. The biggest males would of course have the most impressive growths so in the attached sketch your giant has one of the biggest beards in Earth’s history! What do you think of this idea?
Well, I think it’s awesome. And entirely plausible, for reasons already explained in this post.
“Now, wait,” you may be thinking, “I thought you guys said that sauropod necks weren’t sexually selected.” Actually we made a slightly different point: that the available evidence does not suggest that sexual selection was the primary driver of sauropod neck elongation. But we also acknowledged that biological structures are almost never single-purpose, and although the long necks of sauropods probably evolved to help them gather more food, there is no reason that long necks couldn’t have been co-opted as social billboards. This seems especially likely in Apatosaurus, where the neck length is unremarkable** but the neck fatness is frankly bizarre (and even inspired a Star Wars starfighter!).
I also love the “mobile ecosystem” of birds, other small dinosaurs, and insects riding on this Apatosaurus or following in its train. It’s a useful reminder that we have no real idea what effect millions of sauropods would have on the landscape. But it’s not hard to imagine that most Mesozoic terrestrial ecosystems were sauropod-driven in a thousand cascading and ramifying chains of cause and effect. I’d love to know how that worked. At heart, I’m still a wannabe chrononaut, and all my noodlings on pneumaticity and sauropod nerves and neural spines and so on are just baby steps toward trying to understand sauropod lives. Safari by way of pedantry: tally-ho!
For other speculative apatosaurs, see:
** Assuming we can be blasé about a neck that is more than twice as long (5 m) as a world-record giraffe neck (2.4 m), for garden variety Apatosaurus, or three times that length for the giant Oklahoma Apatosaurus (maybe 7 m).
September 9, 2013
I was at the Oklahoma Museum of Natural History in March to look at their Apatosaurus material, so I got to see the newly-mounted baby apatosaur in the “Clash of the Titans” exhibit (more photos of that exhibit in this post). How much of this is real (i.e., cast from real bones, rather than sculpted)? Most of the vertebral centra, a few of the neural arches, some of the limb girdle bones, and most of the long bones of the limbs. All of the missing elements–skull, neural arches, ribs, appendicular bits–were sculpted by the OMNH head preparator, Kyle Davies. Kyle is one of those frighteningly talented people who, if they don’t have what they need, will just freaking build it from scratch. Over the years he has helped me out a LOT with the OMNH sauropod material–including building a clamshell storage jacket for the referred scapula of Brontomerus so we could photograph it from the lateral side–so it’s about time I gave him some props.
Case in point: this sweet atlas-axis complex that Kyle sculpted for the juvenile Apatosaurus mount.
Most fish, amphibians, and other non-amniote tetrapods only have a single specialized vertebra for attaching to the skull. But amniotes have two: a ring- or doughnut-shaped first cervical vertebra (the atlas) that articulates with the occipital condyle(s) of the skull, and a second cervical vertebra (the axis) that articulates with the atlas and sometimes with the skull as well. Mammals have paired occipital condyles on the backs or bottoms of our skulls, so our skulls rock up and down on the atlas (nodding “yes” motion), and our skull+atlas rotates around a peg of bone on the axis called the odontoid process or dens epistrophei (shaking head “no” motion). As shown in the photos and diagrams below, the dens of the axis is actually part of the atlas that fuses to the second vertebra instead of the first. Also, reptiles, including dinosaurs and birds, tend to have a single ball-shaped occipital condyle that fits into the round socket formed by the atlas, so their “yes” and “no” motions are less segregated by location.
Anyway, the whole shebang is often referred to as the atlas-axis complex, and that’s the reconstructed setup for a baby Apatosaurus in the photo above. In addition to making a dull-colored one for the mount, Kyle made this festive version for the vert paleo teaching collection. Why so polychromatic?
Because in fact he built two: the fully assembled one two photos above, and a completely disassembled one, some of which is shown in this photo (I had to move the bigger bits out of the tray so they wouldn’t block the key card at the back). I originally composed this post as a tutorial. But frankly, since Kyle did all of the heavy lifting of (a) making the thing in the first place, (2) making a color-coded key to it, and (d) giving me permission to post these photos, it would be redundant to walk through every element. So think of this as a self-study rather than a tutorial.
Oh, all right, here’s a labeled version. Note that normally in an adult animal the single piece of bone called the atlas would consist of the paired atlas neural arches (na1) and single atlas intercentrum (ic1), and would probably have a pair of fused cervical ribs (r1). Everything else would be fused together to form the axis, including the atlas pleurocentrum (c1), which forms the odontoid process or dens epistrophei (etymologically the “tooth” of the axis).
Here’s the complete Romer (1956) figure from the key card, with a mammalian atlas-axis complex for comparison. Incidentally, the entire book this is drawn from, Osteology of the Reptiles, is freely available online.
And here’s the complete Gilmore (1936) figure. Sorry for the craptastic scan–amazingly, this one is NOT freely available online as far as I can tell, and Mike and I have been trying to get good scans of the plates for years. Getting back on topic, single-headed atlantal cervical ribs have been found in several sauropods, especially Camarasaurus where several examples are known, so they were probably a regular feature, even though they aren’t always preserved.
Also, as noted in this post, it is odd that in this specimen of Apatosaurus the cervical ribs had not fused to the first two vertebrae, even though they normally do, and despite the fact that the vertebrae had fused to each other, even though they normally don’t. Further demonstration, if any were needed, that sauropod skeletal fusions were wacky.
For comparison to the above images, here is the atlas-axis complex in the synapsid Varanops, from Campione and Reisz (2011: fig. 2C).
Those proatlas thingies are present in some sauropods, but that’s about all I know about them, so I’ll say no more for now.
There is a good overview of the atlas-axis complex with lots of photos of vertebrae of extant animals on this page.
Previous SV-POW! posts dealing with atlantes and axes (that’s right) include:
- A fused atlas and axis in Apatosaurus
- Yet more uninformed noodling on the future of scientific publishing and that kind of thing
- Another mystery: embossed laminae and “unfossae”
- Tutorial 15: the bones of the sauropod skeleton
- Campione, N.E. and Reisz, R.R. 2011. Morphology and evolutionary significance of the atlas−axis complex in varanopid synapsids. Acta Palaeontologica Polonica 56 (4): 739–748.
- Gilmore, C.W. 1936. Osteology of Apatosaurus with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11: 175-300.
- Romer, A.S. 1956. Osteology of the Reptiles. University of Chicago Press, Chicago. 772 pp.