Long-time readers will recall that I’m fascinated by neurocentral joints, and not merely that they exist (although they are pretty cool), but that in some vertebrae they migrate dorsally or ventrally from their typical position (see this and this).

A few years ago I learned that there is a term for the expanded bit of neural arch pedicle that contributes to the centrum in vertebrae with ventrally-migrated neurocentral joints: the bouton, which is French for ‘button’. Here’s an example in the unfused C7 of a subadult sheep. Somebody gifted me a handful of these things a few years ago, and I’ve been meaning to blog about them forever. Many thanks, mysterious benefactor. (I mean, only mysterious to me, because my memory is crap; I’m sure you know who you are, and if you ever read this, feel free to remind me. And thanks for the dead animal parts!)

Guess what? You have these things, too! Or at least you did; if you’re old enough to be reading this, your boutons fused with the rest of the separate bits of your vertebrae a long time ago, between the ages of 2 and 5 (according to Bagnall et al. 1977). Here’s a diagram from Schaefer et al. (2009: p.99) showing the separate centrum and neural arch elements in a thoracic vertebra of a human toddler. So, hey, cool, we all had boutons, just like sheep. And just like some sauropods. (You didn’t think I was going to do a whole OVATOD post without sauropods, did you?)

Here’s our old friend BIBE 45885, an unfused caudal neural arch (or perhaps neural ring) of Alamosaurus, which I’ve been freaking out over for five years now. Those fat bits of neural arch that very nearly close off the neural canal ventrally? Boutons, baby! Big, beautiful boutons. In this photo it looks like the paired boutons meet on the midline, but in fact they merely overlap from this point of view — there is a narrow (<1mm) squiggly gap between them. Given how narrow that gap is, I suspect the two boutons probably would have fused to each other before either of them fused to the centrum, if this particular animal hadn’t died first.

Here’s an unfused dorsal centrum of Giraffatitan, MB.R. 3823, which I yapped about in this post. This vertebra is the spiritual opposite of the Alamosaurus caudal shown above: instead of the neural canal being nearly enclosed by bits of the neural arch wrapping around ventrally, the neural canal is nearly enclosed dorsally by bits of the centrum sticking up on either side and wrapping around dorsally. As with the boutons of the Alamosaurus caudal, the two expanded bits of centrum stuff in this Giraffatitan dorsal approach each other very closely but don’t quite meet; you can fit a piece of paper between them, but not a heck of a lot more. In essence, those “two expanded bits of centrum stuff” are centrum boutons that project up into what I suppose we’ll keep calling a ‘neural arch’ even though it’s neither very neural nor an arch. Or perhaps anti-boutons? With apologies to Gould and Vrba (1982), here we have another missing term in the science of form.

Why do we, and sheep, and prolly lots of other mammals, and some sauropods, have boutons? Presumably to strengthen the neurocentral joints by expanding the joint surface area. I don’t know if anyone has ever tested that — if you do, please let me know in the comments.

Many thanks to Thierra Nalley, who may be the only person I know besides Mike who spends more time thinking about vertebrae than I do, for introducing me to the term ’bouton’ a few years ago. If for some reason you want to corrupt your sensibilities reading about primate vertebrae, you could do a lot worse than checking out Thierra’s papers.

I don’t expect we’ll have a ton of OVATOD posts, in part because there aren’t a heck of a lot of vertebra parts that we haven’t already blogged about. But who knows, maybe Mike will write about prepostepipophyses or something. Stay tuned!


  • Bagnall, K.M., Harris, P.F., and Jones, P.R.M. 1977. A radiographic study of the human fetal spine. 2. The sequence of development of ossification centers in the vertebral column. Journal of Anatomy 124(3): 791–802.
  • Gould, S.J. and Vrba, E.S. 1982. Exaptation—a missing term in the science of form. Paleobiology 8(1): 4-15.
  • Schaefer, M., Black, S., and Scheuer, L. 2009. Juvenile Osteology: A Laboratory and Field Manual. Academic Press, Burlington, MA, 369pp.

P.S. Can we all pitch in and make ’bouton’ the new ‘aglet‘? Please? Please?

I made this for my own amusement, and thought you guys may as well get to benefit from it, too.

Melstrom et al. (2016:figure 4). Pectoral vertebrae of a juvenile specimen of Barosaurus sp. (DINO 2921) from the Upper Jurassic Morrison Formation of Utah, U.S.A., in right lateral view (red-cyan anaglyph made from stereopair).



  • Melstrom, Keegan M., Michael D. D’Emic, Daniel Chure and Jeffrey A. Wilson. 2016. A juvenile sauropod dinosaur from the Late Jurassic of Utah, U.S.A., presents further evidence of an avian style air-sac system. Journal of Vertebrate Paleontology 36(4):e1111898. doi:10.1080/02724634.2016.1111898


I was looking more closely at the turkey skeleton from my recent post, and zeroed in on the last two dorsal (= thoracic) vertebrae. They articulate very well with each other and with the first vertebra of the sacrum, with the centra and zygapophyses both locking in so that there can only have been very little if any movement between them in life. Here they are, in right lateral view:

Last two dorsal (= thoracic) vertebrae of a mature domestic turkey Meleagris gallopavo domesticus, in right lateral view.

Before we move on, it’s worth clicking through to the full-size version of this image and wondering at both the quality of modern phone cameras (a Pixel 3a in this case) and the variety of textures on these little bones. There is smooth, finished bone on the sides of the neural spines; very fine pits and bumps on the zygapophyseal facets where the thin layer of hyaline cartilage attached; rougher texture in the parapophyseal facets where thicker cartilage attached; and very rough texture on the ends of the transverse processes, where there was relatively thick cartilage.

And there is, unsurprisingly in a bird, pneumaticity everywhere. In the more anterior vertebra alone (to the right) the photo shows pneumatic openings (from bottom to top) low on the centrum (below the parapophysis), high on the centrum (below the lateral process),  in the hollow between the lateral process, the posyzyg and the centrum, on the lateral surface of the prezygapophyseal ramus, and on the rear surface of the lateral process. There are others that are obscured in this photo, including on top of the lateral process where it meets the neural spine. Here they are, pointed out for you (with the hidden one shown translucently):

Last two dorsal (= thoracic) vertebrae of a mature domestic turkey Meleagris gallopavo domesticus, in right lateral view. Pneumatic openings on penultimate vertebra highlighted with red lines; obscured opening above lateral process shown as translucent.

OK, that was the B-movie. Now to the main feature. The next photo shows the same two vertebrae, folded away from each other so that we see the anterior face of the posterior vertebra (on the left) and the posterior face of the anterior vertebra (on the right).

Last two dorsal (= thoracic) vertebrae of a mature domestic turkey Meleagris gallopavo domesticus. Left: last dorsal vertebra in anterior view; right: penultimate dorsal vertebra in posterior view.

Again, do click through to see the exquisite detail, especially the complex of pneumatic features on the anterior face of the neural spine of the last dorsal (on the left) and on the posterior face of the left lateral process of the penultimate dorsal (on the right).

And … in the articular facets of the centra?

Seriously, what the heck is going on here? It doesn’t make sense  that there would be pneumatic openings in articular surfaces, because by definition something else (in this case the adjacent vertebra) is abutted hard up against then, so there is no way for a diverticulum to get in. For the same reason, you don’t get vascular foramina in articular surfaces because there is no way to get an artery in there. And there is no hint in these vertebrae of channels along either articular surface that diverticula or arteries could  possibly have laid in.

And yet, there those big openings are. What are they?

I discussed this with Matt, in case it’s Well Known Phenomenon that I’d somehow not heard about but it seems it is not. What we know for sure is that these openings are present, and that they are not mechanical damage inflicted during preparation. So what are they?

What else even is there for them to be? What penetrates bone apart from diverticula and blood vessels? Nerves follow the blood vessels, so it can’t be nerves in the absence of blood vessels.

By the way, there are similar but smaller openings in the posterior face of the last dorsal (the one on the right in the photo), but none anywhere else along the postcervical column: not on the anterior surface of the penultimate dorsal, not on the front or back of the sacrum, and not in any of the other dorsals.

One possibility we considered is that the vertebrae were locked together in life and that a pneumatic space inside the centrum of the last dorsal worked right through into the penultimate one. But that doesn’t work: the openings are not aligned. Also, those in the penultimate dorsal are definitely blind (i.e. they do not connect to deeper internal air-spaces) and those in the last dorsal probably are, too.

We do not know what is going on here.

Help us! Is this kind of thing common in turkeys? Have people seen it in other taxa? Do we know what it is?

Posterior dorsal vertebra of the Upper Cretaceous nanoid saltasaurid LPP-PV-0200. Three-dimensional reconstruction from CT scan in left lateral view (A). Circle and rectangle show sampling planes and the respective thin sections are in (B,C). ce centrum, ns neural spine, pn pneumatopore, poz postzygaphophysis, prz prezygapophysis. Scale bar in (A) 10 cm; in (B,C) 1 cm. Computed tomography data processed with 3D Slicer version 4.10.

Well, this is a very pleasant surprise on the last day of the semester:

Tito Aureliano, Aline M. Ghilardi, Bruno A. Navarro, Marcelo A. Fernandes, Fresia Ricardi-Branco, & Mathew J. Wedel. 2021. Exquisite air sac histological traces in a hyperpneumatized nanoid sauropod dinosaur from South America. Scientific Reports 11: 24207.

You may justly be wondering what I’m doing on a paper on a South American titanosaur. It came about like this:

  • I wrote to Tito Aureliano back in March to congratulate him on his 2019 paper, “Influence of taphonomy on histological evidence for vertebral pneumaticity in an Upper Cretaceous titanosaur from South America”, which I’d just reread, and was impressed by;
  • he told me he was working on a manuscript on saltasaur pneumaticity and would be grateful for my thoughts;
  • I sent him said thoughts, with no strings attached;
  • he asked me if I’d be willing to come on the project as a junior author;
  • I said yes;

and a few months later, here we are.

Dorsal vertebra internal structures of LPP-PV-0200. Reconstructed tomography model in distal (A) and right lateral (B) views illustrating subvertical tangential CT scan slices in false color (1–9). Images show that only a few structures had survived diagenesis which restricted the assessment of the internal architecture to limited spaces. Lighter blue and green indicate lower densities (e.g., pneumatic cavities). Purple and darker blue demonstrate denser structures (e.g., camellate bone). Dashed lines indicate internal plates of bone that sustain radial camellae. ce centrum, cc circumferential chambers, cml camellae, hc-cml ‘honeycomb’ camellae, ns neural spine, pf pneumatic foramen, pn pneumatopore, pacdf parapophyseal-centrodiapophyseal fossa, pocdf postzygapophyseal-centrodiapophyseal fossa, rad radial camellae. Computed tomography data processed with 3D Slicer version 4.10.

My correspondence to Tito basically boiled down to, “All the things you’ve identified in your CT scans are there, but there are also a few more exciting things that you might want to draw attention to” — specifically circumferential and radial camellae near the ends and edges of the centrum, and pneumatic chambers communicating with the neural canal, which were previously only published in Giraffatitan (Schwarz and Fritsch 2006; see Atterholt and Wedel 2018 and this post for more). The internal plates of bone inside the cotyle, which help frame the radial camellae, were first noted by Woodward and Lehman (2009), and discussed in this post.

I can’t think of any reason not to just post the notes I sent to Tito back in March, so here you go:

Wedel suggestions for Aureliano et al Saltasauridae dorsal

I may have more to say about this in the coming days, but at the moment I have two extant dinosaurs — ducks, to be precise — smoking on the grill, and I need to get back to them. The new paper is open access, free to the world (link), so go have fun with it.

UPDATE the next day: here’s another post on the new paper:


In mammals — certainly the most-studied vertebrates — regional differentiation of the vertebral column is distinct and easy to spot. But things aren’t so simple with sauropods. We all know that the neck of any tetrapod is made up of cervical vertebrae, and that the trunk is made up of dorsal vertebrae (subdivided into thoracic and lumbar vertebrae in the case of mammals). But how do we tell whether a given verebra is a posterior cervical or an anterior dorsal?

Here two vertabrae: a dorsal vertebra (D3) and a cervical vertebra (C13) from CM 84, the holotype of Diplodocus carnegii, modified from Hatcher (1901: plates III and VII):

It’s easy to tell these apart, even when as here we have only lateral-view images: the dorsal vertebra is tall, its centrum is short, its neural spine is anteroposteriorly compressed and its parapophysis is up on the dorsal half of the centrum; but the cervical vertebra is relatively low, its centrum is elongated, its neural spine is roughly triangular and its parapophysis hangs down well below the centrum (and has a cervical rib fused to it and the diapophysis).

But things get trickier in the shoulder region because, in sauropods at least, the transition through the last few cervicals to the first few dorsals is gradual — the vertebrae become shorter, taller and broader — and tends to have no very obvious break point. In this respect, they differ from mammals, in which the regional differentiation of the spinal column is more abrupt. (Although even here, things may not be as simple as generally assumed: for example, Gunji and Endo (2016) argued that the 1st thoracic vertebra of the giraffe behaves functionally like an 8th cervical.)

So here are those two vertebrae in context: the sequence D3 D2 D1 C15 C14 C13 in CM 84, the holotype of Diplodocus carnegii, modified from Hatcher (1901: plates III and VII):

Given that the leftmost is obviously a dorsal and the rightmost obviously a cervical, where would you place the break-point?

The most usual definition seems to be that the first dorsal vertebra is the first one that has a free rib, i.e. one not fused to the vertebra: in the illustration above, you can see that the three cervicals on the right all have their cervical ribs fused to their diapophyses and parapophyses, and the three dorsals on the left do not. This definition of the cervical/dorsal distinction seems to be widely assumed, but it is rarely explicitly asserted. (Does anyone know of a paper that lays it out for sauropods, or for dinosaurs more generally?)

But wait!

Hatcher (1903:8) — the same dude — in his Haplocanthosaurus monograph, writes:

The First Dorsal (Plate I., Fig. 1). […] That the vertebra now under consideration was a dorsal is conclusively shown not by the presence of tubercular and capitular rib facets showing that it supported on either side a free rib, for there are in our collections of sauropods, skeletons of other dinosaurs fully adult but, with the posterior cervical, bearing free cervical ribs articulating by both tubercular and capitular facets as do the ribs of the dorsal region. The character in this vertebra distinguishing it as a dorsal is the broadly expanded external border of the anterior branch of the horizontal lamina [i.e. what we would now call the centroprezygapophyseal lamina]. This element has been this modified in this and the succeeding dorsal, no doubt, as is known to be the case in Diplodocus to give greater surface for the attachment of the powerful muscles necessary for the support of the scapula.

Hatcher’s illustrations show this feature, though they don’t make it particularly obvious: here are the last two cervicals and the first dorsal, modified from Hatcher (1903:plate I), with the facet in question highlighted in pink: right lateral view at the top, then anterior, and finally posterior view at the bottom. (The facet is only visible in lateral and anterior views):

Taken at face value, Hatcher’s words here seem to imply that he considers the torso to begin where the scapula first lies alongside the vertebral column. Yet if you go back to the Diplodocus transition earlier in this post, a similar scapular facet is not apparent in the vertebra that he designated D1, and seems to be present only in D2.

Is this scapular-orientation based definition a widespread usage? Can anyone point me to other papers that use it?

Wilson (2002:226) mentions a genetic definition of the cervical/dorsal distinction

Vertebral segment identity may be controlled by a single Hox gene. The cervicodorsal transition in many tetrapods, for instance, appears to be defined by the expression boundary of the Hoxc-6 gene.

But this of course is no use in the case of extinct animals such as sauropods.

So what’s going on here? In 1964, United States Supreme Court Justice Potter Stewart, in describing his threshold test for obscenity, famously said “I shall not today attempt further to define the kinds of material I understand to be embraced within that shorthand description, and perhaps I could never succeed in intelligibly doing so. But I know it when I see it.” Is that all we have for the definition of what makes a vertebra cervicals as opposed to dorsal? We know it when we see it?

Help me out, folks! What should the test for cervical-vs-dorsal be?


  • Gunji, Mego, and Hideki Endo. 2016. Functional cervicothoracic boundary modified by anatomical shifts in the neck of giraffes. Royal Society Open Science 3:150604. doi:10.1098/rsos.150604
  • Hatcher, Jonathan B. 1901. Diplodocus (Marsh): its osteology, taxonomy and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63 and plates I-XIII.
  • Hatcher, J. B. 1903b. Osteology of Haplocanthosaurus with description of a new species, and remarks on the probable habits of the Sauropoda and the age and origin of the Atlantosaurus beds; additional remarks on Diplodocus. Memoirs of the Carnegie Museum 2:1-75 and plates I-VI.
  • Wilson, Jeffrey A. 2002. Sauropod dinosaur phylogeny: critique and cladistic analysis. Zoological Journal of the Linnean Society 136:217-276.

Anatomical features of the neural canal in birds and other dinosaurs. A. MWC 9698, a mid caudal vertebra of Apatosaurus in posterodorsal view. Arrows highlight probable vascular foramina in the ventral floor of the neural canal. B. LACM 97479, a dorsal vertebra of Rhea americana in left anterolateral view. Arrows highlight pneumatic foramina inside the neural canal. C. A hemisected partial synsacrum of a chicken, Gallus domesticus, obtained from a grocery store. Anterior is to the right. The bracket shows the extent of the dorsal recess for the glycogen body, which only spans four vertebrae. Arrows highlight the transverse grooves in the roof of the neural canal for the lumbosacral organ. D. Sagittal (left) and transverse (right) CT slices through the sacrum of a juvenile ostrich, Struthio camelus. The bracket shows the extent of the lumbosacral expansion of the spinal cord. Indentations in the roof of the neural canal house the lumbosacral organ. In contrast to the chicken, the ostrich has a small glycogen body that does not leave a distinct osteological trace. Yellow arrows show the longitudinal troughs in the ventral floor of the neural canal that house the ventral eminences of the spinal cord. Wedel et al. (2021: fig. 4).

This is the second in a series of posts on our new paper about the expanded neural canals in the tail vertebrae of the Snowmass Haplocanthosaurus. I’m not going to talk much about Haplo in this post, though. Instead, I’m going to talk about chickens, and about how you can see a lot of interesting spinal anatomy in a living dinosaur for about two bucks.

You know by now that Academia Letters publishes peer reviews, which is one of the things that drew me to this fairly new journal. More on that in a later post, but in the meantime, the peer reviews for the Haplo paper are on the right sidebar here. I confess, I had a total forehead-slap moment when I read the opening lines of Niels Bonde’s review: 

This paper is interesting, and should be published and discussed by others with interest in dinosaur-bird relations. However, as these publications are also meant for the general public, I would recommend that 2 – 3 illustrations were added of the features mentioned for birds under nos. 3 – 6, because the general public (and many paleontologists) have no ideas about these structures, and what they look like.

The original submission only had figures 1 and 2. And this request is totally fair! If you are going to discuss six alternative hypotheses for some mysterious anatomical structure, it’s just responsible reporting to illustrate those things. That goes double if, as Niels Bonde noted, the anatomy in question is unfamiliar to a lot of people, even many paleontologists. Huxley’s quote after first reading Darwin’s Origin of Species flashed through my head: “How extremely stupid not to have thought of that.”

Slide 21 of my 2014 SVPCA talk on supramedullary diverticula in birds and other dinosaurs, illustrating pneumatic foramina in the roof, walls, and floor of the neural canal.

At the time I read that review, I already had images illustrating five of the six hypotheses. A juvenile ostrich synsacrum that Jessie Atterholt and I had CT scanned gave us three of them all by itself: the lumbosacral expansion of the spinal cord to run the hindlimbs, as in all limbed tetrapods and in some fish with sensitive fins; the transverse channels in the dorsal wall of the neural canal to accommodate the lumbosacral balance organ; and the paired troughs in the floor of the neural canal that house the ventral eminences of the spinal cord (Figure 4D in the image at the top of this post). I had good photos of pneumatic foramina in the walls and floor of the neural canal in a dorsal vertebra of a rhea from my 2014 SVPCA talk (Figure 4B), and some photos of small foramina, presumably for blood vessels rather than air spaces, in the floor of the neural canal in a caudal vertebra of Apatosaurus (Figure 4A).

What I did not have is a photo illustrating the fairly abrupt, dome-shaped space in the sacral neural canal that houses the glycogen body of birds. I mean, I had published images, but I didn’t want to wrestle with trying to get image reproduction rights, or with redrawing the images. Instead, I went to the grocery store to buy some chicken.

I don’t know how universally true this is, but IME in the US when you buy a quartered chicken, the vertebrae are usually nicely hemisected by the band saw that separated the left and right halves of the animals. So you can see the neural canal in both the dorsal and sacral parts of the vertebral column. Here are the hemisected dorsal vertebrae in the breast quarter from a sectioned rotisserie chicken:

That’s just how it came to lie on my plate, but it’s not in anatomical position. Let’s flip it over to sit upright:

And label it:

I could and probably should do a whole post just unpacking this image, but I have other fish to fry today, so I’ll just note a couple of things in passing. The big interspinous ligament is the same one you can see in transverse section in the ostrich dissection photos in this post and this one. Also, the intervertebral joints heading toward the neck, on the left of the image, have much thicker intervertebral cartilage than the more posterior dorsals. That’s because the posterior ones were destined to fuse into a notarium. You can see a diagram and a photograph of a chicken notarium in figures 4 and 5, respectively, here. And finally, the big takeaway here is that the neural canal is normal, just a cylindrical tube to hold the spinal cord.

The thigh quarter usually has the pelvis and the hemisectioned synsacrum attached. Here’s a lateral view of the left half of the pelvis and synsacrum:

And the same thing labeled:

And now flipped around so we can see it in medial view:

And now that image labeled:

And, hey, there are three of our alternative hypotheses on display: the long (many vertebral segments) lumbosacral expansion of the spinal cord, which is reflected in a gradually expanded neural canal in the synsacrum; the shorter, higher dome-shaped recess for the glycogen body; and finally the transverse spaces for the lumbosacral balance organ.

As a refresher, there’s nothing terribly special about the lumbosacral expansion of the spinal cord — you have one, labeled as the ‘lumbar enlargement’ in the above diagram. Where the spinal cord has adjacent limbs to run, it has more neurons, so it gets fatter, so the neural canal gets fatter to accommodate it. The cord itself doesn’t look very expanded in the chicken photo above, but that chicken has been roasted rotisserie-style, and a lot of lipids probably cooked out of the cord during that process. What’s more important is that the neural canal is subtly but unmistakably expanded, over the span of many vertebrae.

The lumbosacral spinal cord of a 3-week-old chick in dorsal view. The big egg-shaped mass in the middle is the glycogen body. Watterson (1949: plate 1).

That’s in contrast to the recess for the glycogen body, which is colored in blue in the chicken photo. Glycogen bodies, like the egg-shaped one in the young chicken in the image immediately above, tend not to go on for many vertebral segments. Instead they balloon up and subside over the space of just 4 or 5 vertebrae, so they leave a different skeletal trace than other soft tissues.

Finally, there are the transverse spaces for the lumbosacral balance organ, which I discussed in this post. Those are the things that look like caterpillar legs sticking up from the sacral endocasts in the above figure from Necker (2006). In life, the spaces are occupied by loops of meningeal membranes, through which cerebrospinal fluid can slosh around, which in turn puts pressure on mechanoreceptive cells at the edge of the spinal cord and gives birds a balance organ in addition to the ones in their heads. In the photo of the cooked chicken, the delicate meninges have mostly fallen apart, leaving behind the empty spaces that they once occupied.

I really liked that chicken synsacrum, and I wanted to use it as part of Figure 4 of the new paper, but it needed a little cleaning, so I simmered it for a couple of hours on low heat (as one does). And it promptly fell apart. At least in the US, most of the chickens that make it to table are quite young and skeletally immature. That particular bird’s synsacrum wasn’t syn-anything, it was just a train of unfused vertebrae that fell apart at the earliest opportunity. I had anticipated that might be an issue, so I’d gotten a lot of chicken, including a whole rotisserie chicken and four thigh quarters from the deli counter at the local supermarket. Happily this fried chicken thigh quarter had a pretty good neural canal:

And it cleaned up nicely:

And with a little cropping, color-tuning, and labeling, it was ready for prime time:

I didn’t label them in the published version, for want of space and a desire not to muddy the waters any further, but the jet-black blobs I have colored in the lower part of that image are the exit holes that let the spinal nerves out of the neural canal so they could go serve the hindlimbs, pelvic viscera, and tail. We have them, too.

At my local grocery store, a fried chicken thigh costs about $1.65 if you get it standalone, or you can buy in bulk and save. You get to eat the chicken, and everything else I’ve done here required only water, heat, soap, and a little time. The point is that if I can do this, you can do this, and if you do, you’ll get to see some really cool anatomy. I almost added, “which most people haven’t seen”, but given how much chicken we eat as a society these days, probably most people’s eyes have fallen on the medial surface of a cooked chicken thigh quarter at one time or another. Better to say, “which most people haven’t noticed”. But now you can. Go have fun. 

Way back in January of 2019, I finished up “Things to Make and Do, Part 25b” with this line: “I have one more thing for you to look for in your bird vertebrae, and that will be the subject of the next installment in this series. Stay tuned!” Here we are, 2.3 years later, and I’ve finally made good. So if there’s a promised post you’ve been waiting for, stick around, we may get to it yet.



FIGURE 7.1. Pneumatic features in dorsal vertebrae of Barapasaurus (A–D), Camarasaurus (E–G), Diplodocus (H–J), and Saltasaurus (K–N). Anterior is to the left; different elements are not to scale. A, A posterior dorsal vertebra of Barapasaurus. The opening of the neural cavity is under the transverse process. B, A midsagittal section through a middorsal vertebra of Barapasaurus showing the neural cavity above the neural canal. C, A transverse section through the posterior dorsal shown in A (position 1). In this vertebra, the neural cavities on either side are separated by a narrow median septum and do not communicate with the neural canal. The centrum bears large, shallow fossae. D, A transverse section through the middorsal shown in B. The neural cavity opens to either side beneath the transverse processes. No bony structures separate the neural cavity from the neural canal. The fossae on the centrum are smaller and deeper than in the previous example. (A–D redrawn from Jain et al. 1979:pl. 101, 102.) E, An anterior dorsal vertebra of Camarasaurus. F, A transverse section through the centrum (E, position 1) showing the large camerae that occupy most of the volume of the centrum. G, a horizontal section (E, position 2). (E–G redrawn from Ostrom and McIntosh 1966:pl. 24.) H, A posterior dorsal vertebra of Diplodocus. (Modified from Gilmore 1932:fig. 2.) I, Transverse sections through the neural spines of other Diplodocus dorsals (similar to H, position 1). The neural spine has no body or central corpus of bone for most of its length. Instead it is composed of intersecting bony laminae. This form of construction is typical for the presacral neural spines of most sauropods outside the clade Somphospondyli. (Modified from Osborn 1899:fig. 4.) J, A horizontal section through a generalized Diplodocus dorsal (similar to H, position 2). This diagram is based on several broken elements and is not intended to represent a specific specimen. The large camerae in the midcentrum connect to several smaller chambers at either end. K, A transverse section through the top of the neural spine of an anterior dorsal vertebra of Saltasaurus (L, position 1). Compare the internal pneumatic chambers in the neural spine of Saltasaurus with the external fossae in the neural spine of Diplodocus shown in J. L, An anterior dorsal vertebra of Saltasaurus. M, A transverse section through the centrum (L, position 2). N, A horizontal section (L, position 3). In most members of the clade Somphospondyli the neural spines and centra are filled with small camellae. (K–N modified from Powell 1992:fig. 16.) [Figure from Wedel 2005.]

Here’s figure 1 from my 2005 book chapter. I tried to cram as much pneumatic sauropod vertebra morphology into one figure as I could. All of the diagrams are traced from pre-existing published images except the horizontal section of the Diplodocus dorsal in J, which is a sort of generalized cross-section that I based on broken centra of camerate vertebrae from several taxa (like the ones shown in this post). One thing that strikes me about this figure, and about most of the CT and other cross-sections that I’ve published or used over the years (example), is that they’re more or less bilaterally symmetrical. 

We’ve talked about asymmetrical vertebrae before, actually going back to the very first post in Xenoposeidon week, when this blog was only a month and a half old. But not as much as I thought. Given how much space asymmetry takes up in my brain, it’s actually weird how little we’ve discussed it.

The fourth sacral centrum of Haplocanthosaurus CM 879, in left and right lateral view (on the left and right, respectively). Note the distinct fossa under the sacral rib attachment on the right, which is absent on the left.

Also, virtually all of our previous coverage of asymmetry has focused on external pneumatic features, like the asymmetric fossae in this sacral of Haplocanthosaurus (featured here), in the tails of Giraffatitan and Apatosaurus (from Wedel and Taylor 2013b), and in the ever-popular holotype of Xenoposeidon. This is true not just on the blog but also in our most recent paper (Taylor and Wedel 2021), which grew out of this post.

Given that cross-sectional asymmetry has barely gotten a look in before now, here are three specimens that show it, presented in ascending levels of weirdness.

First up, a dorsal centrum of Haplocanthosaurus, CM 572. This tracing appeared in Text-fig 8 in my solo prosauropod paper (Wedel 2007), and the CT scout it was traced from is in Fig 6 in my saurischian air-sac paper (Wedel 2009). The section shown here is about 13cm tall dorsoventrally. The pneumatic fossa on the left is comparatively small, shallow, and lacks very distinct overhanging lips of bone. The fossa on the right is about twice as big, it has a more distinct bar of bone forming a ventral lip, and it is separated from the neural canal by a much thinner plate of bone. The fossa on the left is more similar to the condition in dorsal vertebrae of Barapasaurus or juvenile Apatosaurus, where as the one on the right shows a somewhat more extensive and derived degree of pneumatization. The median septum isn’t quite on the midline of the centrum, but it’s pretty stout, which seems to be a consistent feature in presacral vertebrae of Haplocanthosaurus.


Getting weirder. Here’s a section through the mid-centrum of C6 of CM 555, which is probably Brontosaurus parvus. That specific vert has gotten a lot of SV-POW! love over the years: it appears in several posts (like this one, this one, and this one), and in Fig 19 in our neural spine bifurcation paper (Wedel and Taylor 2013a). The section shown here is about 10cm tall, dorsoventrally. In cross-section, it has the classic I-beam configuration for camerate sauropod vertebrae, only the median septum is doing something odd — rather than attaching the midline of the bony floor of the centrum, it’s angled over to the side, to attach to what would normally be the ventral lip of the camera. I suspect that it got this way because the diverticulum on the right either got to the vertebra a little ahead of the one on the left, or just pneumatized the bone faster, because the median septum isn’t just bent, even the vertical bit is displaced to the left of the midline. I also suspect that this condition was able to be maintained because the median septa weren’t that mechanically important in a lot of these vertebrae. We use “I-beam” as a convenient shorthand to describe the shape, but in a metal I-beam the upright is as thick or thicker than the cross bits. In contrast, camerate centra of sauropod vertebrae could be more accurately described as a cylinders or boxes of bone with some holes in the sides. I think the extremely thin median septum is just a sort of developmental leftover from the process of pneumatization.

EDIT 3 days later: John Whitlock reminded me in the comments of Zurriaguz and Alvarez (2014), who looked at asymmetry in the lateral pneumatic foramina in cervical and dorsal vertebrae of titanosaurs, and found that consistent asymmetry along the cervical column was not unusual. They also explicitly hypothesized that the asymmetry was caused by diverticula on one side reaching the vertebrae earlier than diverticula on other other side. I believe they were the first to advance that idea in print (although I should probably take my own advice and scour the historical literature for any earlier instances), and needless to say, I think they’re absolutely correct.

Both of the previous images were traced from CTs, but the next one is traced from a photo of a specimen, OMNH 1882, that was broken transversely through the posterior centrum. To be honest, I’m not entirely certain what critter this vertebra is from. It is too long and the internal structure is too complex for it to be Camarasaurus. I think an apatosaurine identity is unlikely, too, given the proportional length of the surviving chunk of centrum, and the internal structure, which looks very different from CM 555 or any other apatosaur I’ve peered inside. Diplodocus and Brachiosaurus are also known from the Morrison quarries at Black Mesa, in the Oklahoma panhandle, which is where this specimen is from. Of those two, the swoopy ventral margin of the posterior centrum looks more Diplodocus-y than Brachiosaurus-y to me, and the specimen lacks the thick slab of bone that forms the ventral centrum in presacrals of Brachiosaurus and Giraffatitan (see Schwarz and Fritsch 2006: fig. 4, and this post). So on balance I think probably Diplodocus, but I could easily be wrong.

Incidentally, the photo is from 2003, before I knew much about how to properly photograph specimens. I really need to have another look at this specimen, for a lot of reasons.

Whatever taxon the vertebra is from, the internal structure is a wild scene. The median septum is off midline and bent, this time at the top rather than the bottom, the thick ventral rim of the lateral pneumatic foramen is hollow on the right but not on the left, and there are wacky chambers around the neural canal and one in the ventral floor of the centrum. 

I should point out that no-one has ever CT-scanned this specimen, and single slices can be misleading. Maybe the ventral rim of the lateral foramen is hollow just a little anterior or posterior to this slice. Possibly the median septum is more normally configured elsewhere in the centrum. But at least at the break point, this thing is crazy. 

What’s it all mean? Maybe the asymmetry isn’t noise, maybe it’s signal. We know that when bone and pneumatic epithelium get to play together, they tend to make weird stuff. Sometimes that weirdness gets constrained by functional demands, other times not so much. I think it’s very seductive to imagine sauropod vertebrae as these mechanically-optimized, perfect structures, but we have other evidence that that’s not always true (for example). Maybe as long as the articular surfaces, zygapophyses, epipophyses, neural spine tips, and cervical ribs — the mechanically-important bits — ended up in the right places, and the major laminae did a ‘good enough’ job of transmitting forces, the rest of each vertebra could just sorta do whatever. Maybe most of them end up looking more or less the same because of shared development, not because it was so very important that all the holes and flanges were in precisely the same places. That might explain why we occasionally get some really odd verts, like C11 of the Diplodocus carnegii holotype.

That’s all pretty hand-wavy and I haven’t yet thought of a way to test it, but someone probably will sooner or later. In the meantime, I think it’s valuable to just keep documenting the weirdness as we find it.


We’ve noted many times over the years how inconsistent pneumatic features are in sauropod vertebra. Fossae and formamina vary between individuals of the same species, and along the spinal column, and even between the sides of individual vertebrae. Here’s an example that we touched on in Wedel and Taylor (2013), but which is seen in all its glory here:

Taylor and Wedel (2021: Figure 5). Giraffatitan brancai tail MB.R.5000, part of the mounted skeleton at the Museum für Naturkunde Berlin. Caudal vertebrae 24–26 in left lateral view. While caudal 26 has no pneumatic features, caudal 25 has two distinct pneumatic fossae, likely excavated around two distinct vascular foramina carrying an artery and a vein. Caudal 24 is more shallowly excavated than 25, but may also exhibit two separate fossae.

But bone is usually the least variable material in the vertebrate body. Muscles vary more, nerves more again, and blood vessels most of all. So why are the vertebrae of sauropods so much more variable than other bones?

Our new paper, published today (Taylor and Wedel 2021) proposes an answer! Please read it for the details, but here’s the summary:

  • Early in ontogenly, the blood supply to vertebrae comes from arteries that initially served the spinal cord, penetrating the bone of the neural canal.
  • Later in ontegeny, additional arteries penetrate the centra, leaving vascular foramina (small holes carrying blood vessels).
  • This hand-off does not always run to completion, due to the variability of blood vessels.
  • In extant birds, when pneumatic diverticula enter the bone they do so via vascular foramina, alongside blood vessels.
  • The same was probaby true in sauropods.
  • So in vertebrae that got all their blood supply from vascular foramina in the neural canal, diverticula were unable to enter the centra from the outside.
  • So those centra were never pneumatized from the outside, and no externally visible pneumatic cavities were formed.

Somehow that pretty straightforward argument ended up running to eleven pages. I guess that’s what you get when you reference your thoughts thoroughly, illustrate them in detail, and discuss the implications. But the heart of the paper is that little bullet-list.

Taylor and Wedel (2021: Figure 6). Domestic duck Anas platyrhynchos, dorsal vertebrae 2–7 in left lateral view. Note that the two anteriormost vertebrae (D2 and D3) each have a shallow pneumatic fossa penetrated by numerous small foramina.

(What is the relevance of these duck dorsals? You will need to read the discussion in the paper to find out!)

Our choice of publication venue

The world moves fast. It’s strange to think that only eleven years ago my Brachiosaurus revision (Taylor 2009) was in the Journal of Vertebrate Palaeontology, a journal that now feels very retro. Since then, Matt and I have both published several times in PeerJ, which we love. More recently, we’ve been posting preprints of our papers — and indeed I have three papers stalled in peer-review revisions that are all available as preprints (two Taylor and Wedels and a single sole-authored one). But this time we’re pushing on even further into the Shiny Digital Future.

We’ve published at Qeios. (It’s pronounced “chaos”, but the site doesn’t tell you that; I discovered it on Twitter.) If you’ve not heard of it — I was only very vaguely aware of it myself until this evening — it runs on the same model as the better known F1000 Research, with this very important difference: it’s free. Also, it looks rather slicker.

That model is: publish first, then filter. This is the opposite of the traditional scholarly publishing flow where you filter first — by peer reviewers erecting a series of obstacles to getting your work out — and only after negotiating that course to do get to see your work published. At Qeios, you go right ahead and publish: it’s available right off the bat, but clearly marked as awaiting peer-review:

And then it undergoes review. Who reviews it? Anyone! Ideally, of course, people with some expertise in the relevant fields. We can then post any number of revised versions in response to the reviews — each revision having its own DOI and being fixed and permanent.

How will this work out? We don’t know. It is, in part, an experiment. What will make it work — what will impute credibility to our paper — is good, solid reviews. So if you have any relevant expertise, we do invite you to get over there and write a review.

And finally …

Matt noted that I first sent him the link to the Qeios site at 7:44 pm my time. I think that was the first time he’d heard of it. He and I had plenty of back and forth on where to publish this paper before I pushed on and did it at Qeios. And I tweeted that our paper was available for review at 8:44 — one hour exactly after Matt learned that the venue existed. Now here we are at 12:04 my time, three hours and 20 minutes later, and it’s already been viewed 126 times and downloaded 60 times. I think that’s pretty awesome.


  • 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. [PDF]
  • Taylor, Michael P., and Mathew J. Wedel. 2021. Why is vertebral pneumaticity in sauropod dinosaurs so variable? Qeios 1G6J3Q. doi: 10.32388/1G6J3Q [PDF]
  • Wedel, Mathew J., and Michael P. Taylor 2013b. 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]

You! Shall not! Pass!

August 22, 2020

OK, technically this is MB.R.3822, a dorsal vertebra of Giraffatitan brancai formerly known as HMN Ar1, in posterior view, rendered from a 3D scan provided by Heinrich Mallison.

But you can’t tell me that when you look at that you don’t see Gandalf shouting at a balrog.

Long before Matt and others were CT-scanning sauropod vertebrae to understand their internal structure, Werner Janensch was doing it the old-fashioned way. I’ve been going through old photos that I took at the Museum für Naturkunde Berlin back in 2005, and I stumbled across this dorsal centrum:

Dorsal vertebra centum of ?Giraffatitan in ventral view, with anterior to top.

You can see a transverse crack running across it, and sure enough the front and back are actually broken apart. Here there are:

The same dorsal vertebral centrum of ?Giraffatitan, bisected transversely in two halves. Left: anterior half in posterior view; right: posterior half in anterior view. I had to balance the anterior half on my shoe to keep it oriented corrrectly for the photo.

This does a beautiful job of showing the large lateral foramina penetrating into the body of the centrum and ramifying further into the bone, leaving only a thin midline septum.

But students of the classics will recognise this bone immediately as the one that Janensch (1947:abb. 2) illustrated the posterior half of in his big pneumaticity paper:

It’s a very strange feeling, when browsing in a collection, to come across a vertebra that you know from the literature. As I’ve remarked to Matt, it’s a bit like running into, say, Cameron Diaz in the corner shop.


  • Janensch, W. 1947. Pneumatizitat bei Wirbeln von Sauropoden
    und anderen Saurischien. Palaeontographica, supplement