Tutorial 36: the atlas-axis complex in sauropods
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.
UPDATE in December, 2019: oh heck with it, I’m very belatedly renaming this a tutorial, so I can tag on a follow-up post as Tutorial 36b and curate this where it belongs, on our Tutorials page. The URL will stay the same, like a digital fossil.
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
References
- 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.
September 9, 2013 at 5:56 am
By deduction from contextual clues, I am compelled to conjecture that these osteoliths constitute the neck-bone that’s connected to the head-bone.
September 9, 2013 at 6:06 am
Ah, er, I seem to have neglected to mention what the atlas and axis are. *chagrin* You got it, they are cervicals 1 and 2. I will amend the post.
September 9, 2013 at 12:57 pm
@Nathan Myers
Ah, yes, but could they raise them above the horizontal position!
September 9, 2013 at 1:23 pm
Good post! it is about what i´m working right now, and will be helpful. I love the model made by Kyle Davies too.
September 9, 2013 at 5:51 pm
[…] composites of many specimens, or have missing parts filled in with casts or sculpted elements (see this SVPOW post on Kyle Davies, who sculpts bone replicas for OMNH). Probably the most important function of a mount is to present […]
September 9, 2013 at 8:05 pm
@anon: Trick question: they would have to get the neck into a horizontal position, first.
September 12, 2013 at 7:53 pm
That looks not mobile.
September 13, 2013 at 4:22 am
Right. It’s not. The presence of fairly long cervical ribs* on the atlas and the fusion of the atlas and axis in CM 3018 are evidence that almost no motion could take place at the atlas-axis joint, unlike mammals, where there is a lot of rotation between atlas and axis. But whereas mammals can only rock our heads up and down on our dual occipital condyles, dinos had more of a ball-and-socket setup between the single occipital condyle the bony cup formed by the atlas. So the lack of movement between atlas and axis was not a problem.
* Mammalian cervicals, including the atlas, have bicipital (two-headed) cervical ribs, but they’re super-short and they always fuse to the vertebrae, so we refer to them as transverse processes. But developmentally they are cervical ribs, the same as those in birds and sauropods.
September 14, 2013 at 12:06 am
[…] Sauropod neck bones are cool. […]
February 14, 2014 at 9:21 am
[…] (How could Lull have been unsure whether the most anterior preserved cervical was the second or third? C2 in sauropods, as in most animals, is radically different from the subsequent cervicals. He does go on to say that only the centrum of the most anterior vertebra is preserved, but the axis has a distinct anterior central articulation.) […]
April 18, 2014 at 8:45 pm
[…] objects in the universe. Also, an appropriately huge thank-you to preparator Kyle Davies (of apatosaur-sculpting fame), collections manager Jen Larsen, and Andrew Thomas again for help with wrassling those verts […]
August 19, 2015 at 6:20 am
[…] was preserved in perfect articulation, even the hyoid apparatus, terminal phalanges, proatlas, and atlas cervical ribs. The skull was a bit disarticulated – half of the rostrum had floated out of position, and […]
February 27, 2016 at 8:43 am
[…] this shot from behind, you get a better look at the baby apatosaur standing under the big one, and it hints at a far more likely target for Saurophaganax and other […]
December 21, 2016 at 6:51 pm
[…] and general sculpting/molding/casting sorceror. You’ve seen his work on the baby apatosaur in this post. I have casts of everything shown here – original fossil, fossil-plus-prosthesis, and […]
December 22, 2017 at 7:06 am
[…] year Santaposeidon comes to you courtesy of OMNH vert paleo head preparator and 20th-level fossil conservation wizard Kyle Davies, who took the photo, composed the card, and gave me kind permission to share it here. […]
January 2, 2018 at 12:23 am
[…] blogged a lot about the giant – and tiny – apatosaurines from the Morrison Formation of Oklahoma, and just once on Saurophaganax. But […]
December 8, 2019 at 1:44 am
[…] beyond my capacity, but we can roll with what I have. Before we go on, you might want to revisit Tutorial 36 to brush up on the general parts of the atlas-axis […]
November 23, 2022 at 8:34 am
[…] In this picture, the atlas seems to be pretty much fused onto the axis, as seen in Gilmore (1936: figure 6) which Matt helpfully reproduced in Tutorial 36. […]