Last Thursday I gave a public lecture for the No Man’s Land Historical Society in the Oklahoma Panhandle, titled “Oklahoma’s Jurassic Giants: the Dinosaurs of Black Mesa”. It’s now on YouTube, on the No Man’s Land Museum’s channel.

There’s a point I want to make here, that I also made in the talk: we can’t predict the value of natural history collections. The first sauropod vertebrae that Rich Cifelli and Kent Sanders and I CT scanned back in the spring of 1998 belonged to what would become Sauroposeidon, but most of the ones we scanned after that were Morrison specimens collected by J. Willis Stovall’s crews from the Oklahoma Panhandle between 1934 and 1941. Those scans formed the core of the pneumaticity research that fleshed out the Sauroposeidon papers (Wedel et al. 2000a, b), and was more fully developed in my Master’s thesis and the papers that came out of that (Wedel 2003a, b).

OMNH 1094, a mid-cervical vertebra of Brontosaurus in right lateral view. If you’ve seen one of my talks or my first few papers, you’ve seen this vert. I just realized that I have almost all the photos I need to do a proper multi-view; stand by for a future post on that.

So the foundation of my career was built in large part from collections that had been made 60 years earlier, decades before CT was invented. I’ll also note here that Xenoposeidon — Mike’s fourth paper (Taylor and Naish 2007), but the one which really launched his career as a morphologist — is based on a specimen collected in the 1890s. Natural history collections are not only resources for making comparisons, but also the engines of future discovery, and building and maintaining them is one of the most significant contributions to science that we can make.

I thank a bunch of folks at the end of the talk, but I especially want to thank Brian Engh for the use of his art, and Anne Weil for inviting me to collaborate on the sauropod material from the Homestead Quarry. Looking forward to more adventures!


Morphological variation in paramedullary airways; yellow = spinal cord, green = diverticula. The spectrum of variation is discretized into four groups: i, branches of intertransverse diverticula contact spinal cord at intervertebral joints; ii, branches of intertransverse diverticula extend partially into the vertebral canal, but remain discontinuous; iii, paramedullary diverticula form sets of tubes that are continuous through vertebral canals of at least two consecutive vertebrae; iv, continuous paramedullary diverticula anastomose with supravertebral diverticula. Each variant is depicted diagrammatically (A, dorsal view; B, E, H, & K, transverse view) and shown in two CT scans; images in each column correspond to the same morphology. Morphology i: C, cormorant; D, scrub jay. Morphology ii: F, bushtit; G, common murre. Morphology iii: I, red-tailed hawk; J, black-crowned night heron. Morphology iv: L, M, pelican. (Atterholt and Wedel 2022: figure 5)

New paper out:

Atterholt, Jessie, and Wedel, Mathew J. 2022. A computed tomography-based survey of paramedullary diverticula in extant Aves. The Anatomical Record, 1– 22.

Quick aside, which will soon be of historical interest only: so far, only the accepted-but-unformatted manuscript is available, with the final, fully-formatted ‘version of record’ due along at some point in the future. We’re not sure when that will be — could be next week, could be months from now — which is why I’m following my standard procedure and yapping about the new paper now. This has paid off in the past, when papers that were only available in accepted ms form were read and cited before the final version was published. UPDATE on April 9: the formatted version of record is out now, as an open-access article with a CC-BY license, and I swapped it for the ‘accepted ms’ version in the links above and at the end of this post.

This paper has had a weirdly drawn-out gestation. Jessie and I hatched the idea of it way back in 2017, when we were teaching in the summer anatomy course together. I learned that Jessie had a big war chest of CTs of dead birds, and I’d been obsessed with supramedullary diverticula in birds and sauropods for some time already (e.g., an SVPCA talk: Wedel et al. 2014). There were detailed published descriptions of the supramedullary diverticula in a handful of species — namely chickens, turkeys, and pigeons — but no broad survey of those diverticula across living birds. Jessie had the CT scans to do that big survey, which we got rolling on right away. She presented our preliminary results at SVPCA in 2018 (Atterholt and Wedel 2018), and by rights the paper should have been along shortly thereafter. More on that in a sec.

One thing that may seem odd: we use the term ‘paramedullary diverticula’ instead of the more familiar and established ‘supramedullary diverticula’. That’s because these diverticula are not always dorsal to the spinal cord; sometimes they’re lateral, sometimes they’re ventral, and sometimes they completely surround the spinal cord, like an inflated cuff. So we decided that the term ‘paramedullary’, or ‘next to the spinal cord’, was more accurate than ‘supramedullary’, or ‘above the spinal cord’, for describing this class of diverticula.

Observed variation in the shape, arrangement, and orientation of paramedullary diverticula relative to the spinal cord; yellow = spinal cord, green = diverticula. A, paired diverticula dorsal to spinal cord in an ostrich. B, paired diverticula lateral to spinal cord in a bushtit. C, paired diverticula ventral to spinal cord in a violet turaco. D, three diverticula dorsal to spinal cord in an ostrich. E, four diverticula dorsal to spinal cord in an eclectus parrot. F, single, c-shaped diverticulum dorsal to spinal cord in an ostrich. G, diverticula completely surrounding spinal cord and pneumatizing vertebra in a violet turaco. H, no paramedullary diverticula present in a Pacific loon. I, diverticula completely surrounding spinal cord in a pelican. (Atterholt and Wedel 2022: figure 6)

I will have more to say about the science in other posts, and you can get the scientific backstory in this post and this one and the abstracts cited above and linked below. The rest of this post is mostly about me, so if you stick around, buckle up for some advanced navel-gazing.

There’s no one reason why this paper didn’t come out sooner. In short, I hit a wall. We went through a curriculum change at work, and suddenly the annual schedule that I’d relied on for a decade was completely upended. I had some unexpected challenges in my personal life. But the biggest problem was that my attitude toward research and writing had changed, for the worse.

When I was fresh out of grad school I had this kinda snotty attitude that my research was MINE, and wherever I was teaching was just, like, a paycheck, man, but they don’t own me, or my research. And as my teaching and committee responsibilities ramped up I still felt like research and writing was something I did for myself, and that my mission was to steal however many hours I could away from the “day-job work” to get done the things that I really wanted to do. Like a guerilla insurgency. In retrospect, it was a pretty good attitude for getting stuff done.

But somewhere along the way, I stopped thinking about research as something that belonged to me, something that I did for myself, and started thinking about it as part of my job. (This also maybe is not so flattering in what it reveals about how I think, or at least thought, about my job.) Instead of using my research time as a source of energy and a wellspring of satisfaction and positivity, I starting thinking of it only as a sink. And it happened so insidiously that I didn’t even realize it. My productivity plummeted, and I didn’t understand why. I was restless and depressed, and I didn’t understand that either. At the level of my superficial thoughts I still wanted to get research done, but my subconscious was turned off to it, so I just spun my wheels.

Then the pandemic hit. I’d always been a pretty optimistic, upbeat person, but I found myself just staring off into space franticizing about all the horrible things going on in the world, or staying up too late doom-scrolling the news. I slept too little, and poorly, and by the end of 2020 I felt worn down to a nub.

Osteological evidence of paramedullary diverticula. A, pocked texturing inside the vertebral canal of a pelican (LACM 86262). B, pneumatic foramen on the roof of the vertebral canal of an albatross (Phoebastria nigripes, LACM 115139). C, pneumatic foramina in the floor of the vertebral canal of an ostrich (Struthio camelus, LACM 116205). D, deep pneumatic fossae in the roof of the vertebral canal of an Eastern moa (Emeus sp., LACM unnumbered). (Atterholt and Wedel 2022: figure 7)

Then a series of positive things happened:

  • I received a long, heartfelt email from Jessie (fittingly!), asking after me and laying out a plan for getting the paper done and out. It was the kick I needed to look inside and start picking myself apart, to figure out what the heck was going on. Much of this post is cribbed from my reply to her.
  • I got a little break from lecturing in the spring of 2021, and that gave me the space to get a couple of things finished and submitted — the pneumatic variation paper with Mike in January (Taylor and Wedel 2021), and the Haplocanthosaurus neural canal paper, which was submitted even earlier in January, although it came out much later (Wedel et al. 2021; more on that publication delay in a future post).
  • Finally, I had young, energetic coauthors who were moving projects forward that required modest levels of effort from me, but which paid off with highly visible publications that I’m proud to be an author on, including the saltasaur pneumaticity paper (Aureliano et al. 2021) and the ‘Sauro-Throat’ paper (Woodruff et al. 2022).

It’s impossible to overstate how energizing it was to get new things done and out, and how much it helped to have collaborators who were putting wins on the board even when I was otherwise occupied. One of those collaborators was Jessie, who just kept pushing this thing forward — and, sometimes, pushing me forward — until it was done. So the paper you can read today is a testament not only to her acumen as a morphologist, but also to her tenacity as a scholar, and as a friend.

The part of the paper I’m happiest about is the “Conclusions and Directions for Future Research”, which points the way toward a LOT of further studies that need to be done, both on extant birds and on fossil archosaurs, ranging from bone histology to functional morphology to macroevolution. As we wrote in the concluding sentence of the paper, “We hope that this study serves as a foundation and an enticement for further studies of this most unusual anatomical system, in both extinct and extant archosaurs.”

I can’t wait to see what comes next.


Science doesn’t always get done in the right order.

In the course of the research for my paper with Mike this past spring, “Why is vertebral pneumaticity in sauropod dinosaur so variable?”, published in Qeios in January, I had a couple of epiphanies. The first was that I had collated enough information to map the sites at which arteries and veins enter and exit the vertebrae in most tetrapods. The second was that, having done that, I’d also made a map of (almost) all the places that diverticula enter the vertebrae to pneumatize them. This is obviously related to the thesis we laid out in that paper, that postcranial skeletal pneumaticity is so variable because pneumatic diverticula follow pre-existing blood vessels as they develop, and blood vessels themselves are notoriously variable. In fact, if you had to summarize that thesis in one diagram, it would probably look like the one above, which I drew by hand in my research notebook in early March.

Only that’s not quite correct. I didn’t have those epiphanies “in the course of the research”, I had them after the pneumatic variation paper was done and published. And at the time they felt less like epiphanies and more like a series of “Holy crap” realizations:

  1. Holy crap, that diagram would have been really helpful when we were writing the pneumatic variation paper, since it establishes, almost tautologically, that diverticula invade vertebrae where blood vessels already have. In a rational world, Mike and I would have done this project first, and the pneumatic variation paper would have stood on its shoulders.
  2. Holy crap, how have I been working on vertebral pneumaticity for more than two decades without ever creating a map of all the places a vertebra can be pneumatized, or even realizing that such a map would be useful?
  3. Holy crap, how have I been working on dinosaur bones — and specifically their associated soft tissues — for more than two decades without wondering exactly how the blood was getting into and out of each bone? 

Arguably, not only should Mike and I have done this project first, I should have taken a stab at it way back when I was working on my Master’s thesis. Better late than never, I guess.

I used a sauropod caudal as my vertebral archetype because it has all the bits a tetrapod vertebra can have, including the hemal arch or chevron. This was important, because Zurriaguz et al. (2017) demonstrated that the chevrons are pneumatic in some titanosaurs. 


For the actual presentation I redrew the vessels on top of a scan of a Camarasaurus caudal from Marsh, which Mike found and cleaned up (modified from Marsh 1896: plate 34, part 4, and plate 39, part 3c). 

We deliberately used an unfused caudal to emphasize that ‘ribs’ — technically, costal elements — are present, they just fuse to the neural arch and centrum rather than remaining separate, mobile elements like dorsal ribs.

Anyway, I’m yapping about this now because this project is rolling: Mike and I submitted an abstract on it for the 3rd Palaeontological Virtual Congress, and a short slideshow on the project is now up at the 3PVC site for attendees to look at and comment on. The congress started last Wednesday and runs through Dec. 15, after which I’m sure we’ll submit the abstract and slide deck somewhere as a preprint, and then turn it into a paper as quickly as possible.

I’ll probably have more to say on this in a day or so, but for now the comment field is open, and your thoughts are welcome.



FMNH P13018 with me for scale. Photo by Holly Woodward.

Some of the Burpee Museum folks and PaleoFest speakers visited the Field Museum of Natural History in Chicago after the 2020 ‘Fest. I hadn’t been there since 2012, and a lot had changed. More on that in future posts, maybe. Here I am with FMNH 13018, a right femur referred by von Huene (1929) to Argyrosaurus superbus (note, though, that Mannion and Otero 2012 considered this specimen to be Titanosauria indet., hence the hedge in the title of the post). It’s 211cm long, which is pretty darn big but still well short of the record.

Speaking of the record, here’s a list of the largest sauropod femora (as always, updates in the comments are welcome!):

  1. 250cm – Argentinosaurus huinculensis, MLP-DP 46-VIII-21-3 (estimated when complete)
  2. 238cm – Patagotitan mayorum, MPEF-3399/44
  3. 236cm – Patagotitan mayorum, MPEF-PV 3400/27
  4. 235cm – Patagotitan mayorum, MPEF-PV 3400/27
  5. 235cm – “Antarctosaurus” giganteus, MLP 26-316
  6. 214cm – Giraffatitan brancai, XV1
  7. 211cm – cf. Argyrosaurus superbus, FMNH P13018
  8. 203cm – Brachiosaurus altithorax, FMNH P25107
  9. 200cm – Ruyangosaurus giganteus, 41HIII -0002 (estimated when complete)
  10. 191cm – Dreadnoughtus schrani, MPM-PV 1156

The list is necessarily incomplete, because we have no preserved femora for Puertasaurus, Notocolossus, Futalognkosaurus, or the largest individuals of Sauroposeidon and Alamosaurus, all of which probably had femora in the 210-250cm range. For that matter, most elements of the giant Oklahoma apatosaurine are 25%-33% larger than the equivalent bones in CM 3018, which implies a femur length of 223-237cm (scaled up from the 178.5cm femur of CM 3018). I’m deliberately not dealing with Maraapunisaurus or horrifying hypothetical barosaurs here.

In any case, it’s still a prodigious bone, and well worth spending a moment with the next time you’re at the Field Musuem.


  • Mannion, P.D. and Otero, A., 2012. A reappraisal of the Late Cretaceous Argentinean sauropod dinosaur Argyrosaurus superbus, with a description of a new titanosaur genus. Journal of Vertebrate Paleontology, 32(3):614-638.
  • Von Huene, F. 1929. Los saurisquios y ornitisquios del Creta´ceo Argentino. Anales del Museo de La Plata 3:1–196.

Here’s a pretty cool image: Plate 7 from Lull (1919), showing the partial skeleton of Barosaurus YPM 429 (above), compared to the much more complete skeleton of Diplodocus CM84/94 (below).

I’ve been pretty familiar with that Barosaurus skeleton diagram since I was about 9 years old, because it’s in Donald Glut’s New Dinosaur Dictionary, which I’ve written about here before. In particular, I like that Lull was scrupulous about drawing in the lateral pneumatic cavities in the caudal vertebrae. It’s pretty common in Diplodocus for the tail to be pneumatized out to somewhere between caudal 15 and 19, and the same is true in Barosaurus. I’m not just relying on the figure–Lull was also good about saying explicitly what was going on with the pneumatization in the centrum of each vertebra.

I returned to this image as an adult doing research on sauropod pneumaticity, and I read big swaths of Lull (1919), but never the bit about the sacrum. Why would I? The sacrum of YPM 429 is pretty scrappy, and I was mostly interested in the big honkin’ cervicals, and in learning how to distinguish bones of Barosaurus and Diplodocus. I always assumed that the sacrum of Barosaurus was pneumatized right the way through.

Only, er, it ain’t. As I just discovered.

Lull (1919: p. 22):

See that second sentence? “The central fragment is extremely massive, with no adaptation for lightening the weight appreciable in the portion preserved.” That’s old-timey talk for, “the chunk of centrum has no pneumatic openings or cavities”. Which is kind of a big deal, because:

…a gap of one or more apneumatic vertebrae with pneumatic vertebrae on either side constitutes a pneumatic hiatus. Why that’s a big deal is explained in this post.

If I had read this in the early 2000s, I would have flipped out. I did flip out when I discovered what seemed to be a pneumatic hiatus at the base of the tail in Haplocanthosaurus. Just that possibility sent me scurrying off to the Carnegie Museum to investigate, and precipitated both a dissertation chapter, later published as Wedel (2009), and an enduring fascination with Haplocanthosaurus. If I’d been reading Lull instead of Hatcher, my air sac paper would have been about Barosaurus, probably, and I wouldn’t have known enough about Haplo to get interested in the other specimens, which would have been a real shame.

A pneumatic hiatus in Barosaurus would have been big news in 2009. In 2021, it’s still nice, but not groundbreaking. The groundbreaking pneumatic hiatuses in Barosaurus were described in two different juvenile skeletons by Melstrom et al. (2016) and Hanik et al. (2017). Those were both mid-thoracic hiatuses, which probably separated the pneumatization domains of the cervical air sacs anteriorly and the abdominal air sacs posteriorly. A mid-sacral hiatus in YPM 429 is probably within the domain of the abdominal air sac, just like the hiatus in sacral 5 of CM 879 that I described in my 2009 paper. It’s still exciting, in that it shows that there were abdominal air sacs, and they were separate from the lungs and cervical air sacs, but this example in YPM 429 is now third in line in terms of priority, just within this one genus. Which is why I’m telling the world with a blog post, instead of hopping on a plane (or, er, planning a very long road trip) to New Haven. I’ll check on YPM 429 the next time I’m out there, but the specifics will keep for now.


  • Hanik, Gina M., Matthew C. Lamanna and John A. Whitlock. 2017. A juvenile specimen of Barosaurus Marsh, 1890 (Sauropoda: Diplodocidae) from the Upper Jurassic Morrison Formation of Dinosaur National Monument, Utah, USA. Annals of Carnegie Museum 84(3):253–263.
  • Lull, R.S. 1919. The sauropod dinosaur Barosaurus Marsh. Memoirs of the Connecticut Academy of Arts and Sciences 6:1-42.
  • Melstrom, Keegan M., Michael D. D’Emic, Daniel Chure and Jeffrey A. Wilson. 2016. A juvenile sauropod dinosaur from the Late Jurassic of Utah, USA, presents further evidence of an avian style air-sac system. Journal of Vertebrate Paleontology 36(4):e1111898. doi:10.1080/02724634.2016.1111898
  • Wedel, M.J. 2009. Evidence for bird-like air sacs in saurischian dinosaurs. Journal of Experimental Zoology 311A:611-628.

A. Recovered skeletal elements of Haplocanthosaurus specimen MWC 8028. B. Caudal vertebra 3 in right lateral view. C. The same vertebra in posterior view. Lines show the location of sections for D and E. D. Midsagittal CT slice. The arrow indicates the ventral expansion of the neural canal into the centrum. E. Horizontal CT slice at the level of the neural arch pedicles, with anterior toward the top. Arrows indicate the lateral expansions of the neural canal into the pedicles. B-E are shown at the same scale. Wedel et al. (2021: fig. 1).

New paper out today:

Wedel, Mathew; Atterholt, Jessie; Dooley, Jr., Alton C.; Farooq, Saad; Macalino, Jeff; Nalley, Thierra K.; Wisser, Gary; and Yasmer, John. 2021. Expanded neural canals in the caudal vertebrae of a specimen of Haplocanthosaurus. Academia Letters, Article 911, 10pp. DOI: 10.20935/AL911 (link)

The paper is new, but the findings aren’t, particularly. They’re essentially identical to what we reported in our 1st Paleo Virtual Conference slide deck and preprint, and in the “Tiny Titan” exhibit at the Western Science Center, just finally out in a peer-reviewed journal, with better figures. The paper is open access and free to the world, and it’s short, about 1600 words, so this recap will be short, too.

A. Photograph of a 3D-printed model of the first three caudal vertebrae of Haplocanthosaurus specimen MWC 8028, including endocasts of the neural canal (yellow) and intervertebral joints (blue), in right lateral view, and with the neural canal horizontal. B. Diagram of the same vertebrae in midsagittal section, emphasizing the volumes of the neural canal (yellow) and intervertebral joint spaces (blue). Anterior is to the right. Wedel et al. (2021: fig. 2).

John Foster and I described Museum of Western Colorado (MWC) specimen 8028, a partial skeleton of Haplocanthosaurus from Snowmass, Colorado, in late 2014. One weird thing about that specimen (although not the only weird thing) is that the neural canals of the tail vertebrae are bizarrely expanded. In most vertebrae of most critters, the neural canal is a cylindrical tunnel, but in these vertebrae the neural canals are more like spherical vacuities.

John and I didn’t know what to make of that back in 2014. But a few years later I started working with Jessie Atterholt on bird anatomy, which led me to do a little project on the whole freaking zoo of weird stuff that birds and other dinosaurs do with their neural canals, which led to the 1PVC presentation, which led to this. 

Caudal vertebra 3 of Haplocanthosaurus specimen MWC 8028 in left posterolateral (A), posterior (B), and right posterolateral (C) views, with close-ups (D and E). In A and B, a paintbrush is inserted into one of the lateral recesses, showing that the neural canal is wider internally than at either end. Wedel et al. (2021: fig. 3).

Of course there will be more posts and more yapping, as signaled by the ‘Part 1’ in the post title. Although I am extremely satisfied with the streamlined, 1600-word missile of information and reasoning that just dropped, there are parts that I want to unpack, that haven’t been unpacked before. But the paper launched at midnight-thirty, Pacific Daylight Time, I’m up way too late finishing this first post, and I reckon the rest will keep for a few hours at least.

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).

I have a ton of people to thank. John Foster, obviously, for initiating the line of research that led here. Julia McHugh for access to the MWC collections, and for being an excellent sounding board regarding the Morrison Formation, sauropod dinosaurs, and crafting ambitious but tractable research projects. Anne Weil for helping me be methodical in thinking through the logic of the paper, and Mike Taylor for helping me get it polished. Niels Bonde, Steven Jasinski, and David Martill for constructive reviews, which were published alongside the paper. We couldn’t take all of their suggestions because of space limitations, but figures 3 and 4 were born because they asked for them, and that’s not a small thing. Vicki and London Wedel for putting up with me at various points in this project, especially in the last few days as I’ve been going bonkers correcting page proofs. And finally, because I’m the one writing this blog post, my coauthors: Jessie Atterholt, Alton Dooley, Saad Farooq, Jeff Macalino, Thierra Nalley, Gary Wisser, and John Yasmer, for their contributions and for their patience during the unusually long gestation of this very short paper.

More to say about all that in the future. For now, yay, new paper. Have fun with it. Here’s the link again.


Back in 2017, I showed the world 83.33% of my collection of sauropod-themed mugs. Time passes, and I have lost some of them and gained some more. The tally now stands at eight, and here they are:

My missing Brontomerus mug never did turn up. In the mean time, I have also lost or maybe broken the Sauroposeidon mug, the old black-and-white Archbishop mug, and the single-view Xenoposeidon mug. The dissertation mug still survives, but has faded into total illegibility, so I don’t count it any more.

On the more positive side, the sexual selection mug — second from the right in the old photo, and bottom left in the new one — survives, in fact the only one to have done so. All the others are new acquisitions. Let’s take a look:

Back row, left to right:

  1. The new, improved Archbishop dorsals A and B mug. Unlike the original, this is in glorious colour, and rearranges the elements to show anterior view on the front, and left and right lateral on the sides.
  2. The new, improved Xenoposeidon mug. It’s laid out the same way with the anterior view on the front and left and right lateral views on the sides.
  3. One that Fiona made for my birthday, showing one of the publicity photos from the original Xenoposeidon description: the one of which a newspaper columnist wrote “I wish my husband looked at me the way he looks at this bone”.
  4. A mug made by Mark Witton, which I saw at TetZooCon 2019 and made him an offer for. It shows his own Diplodocus artwork, an update of an earlier piece that he did for Matt, Darren and me to publicise our 2009 paper on sauropod neck posture. (Details here.)

Front row, left to right:

  1. The sole survivor, showing the introductory here’s-what-sauropod-necks-are-like illustration from our 2011 paper on why those necks were not sexually selected.
  2. The sauropod neck gallery used as Figure 3 in my and Matt’s 2013 PeerJ paper “why giraffes have short necks”.
  3. One of the world’s few caudal pneumaticity mugs, using all the illustrations from Matt’s and my 2013 paper, and inspired by the freakily consistent colour palette of those illustrations.
  4. This one needs a bit of explaining. See below.

For reasons that no-one — least of all he — understands, my youngest son bought a pair of Dawn French mugs as a birthday or Christmas present for Fiona. (No-one in our family is particularly a fan, it was one of those random things.) Since then, he has given her five or six more identical mugs.

Because I do not like these, I insist that they hang on one mug tree, and the sauropod mugs on another. It was to break down this mug apartheid that our eldest made for us this final mug, which shows both Dawn French and a reconstruction of the Xenoposeidon vertebra (from my 2018 paper). Where does it live? Usually, it sits on the shelf between the two mug trees.

So this is how things stand. (I drink a lot of tea, so these mugs all see plenty of action.) I really should make myself a new Brontomerus mug, and perhaps a pneumatic variation one.


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.


Figure 3. BIBE 45854, articulated series of nine mid and posterior cervical vertebrae of a large, osteologically mature Alamosaurus sanjuanensis. Series is estimated to represent the sixth to fourteenth cervical vertebrae. A, composite photo-mosaic of the cervical series in right lateral view; identification of each vertebra indicated by C6 to C14, respectively. B, line drawing based on the photo-mosaic in A. C, line drawing in B with labels shown and vertebral fossae indicated by solid grey fill; cross-hatching represents broken bone surfaces and reconstructive material. Abbreviations: C, cervical vertebra; cdf, centrodiapophyseal fossa; clf, centrum lateral fossa; pocdf, postzygapophyseal centrodiapophyseal fossa; prcdf, prezygapophyseal centrodiapophyseal fossa; prcdf1, dorsal prezygapophyseal centrodiapophyseal fossa; prcdf2, ventral prezygapophyseal centrodiapophyseal fossa; sdf, spinodiapophyseal fossa; spof, spinopostzygapophyseal fossa; sprf, spinoprezygapophyseal fossa. (Tykoski and Fiorillo 2016)

Have you been reading Justin Tweet’s series, “Your Friends the Titanosaurs“, at his awesomely-named blog, Equatorial Minnesota? If not, get on it. He’s been running the series since June, 2018, so this notice is only somewhat grotesquely overdue. The latest installment, on Alamosaurus from Texas and Mexico, is phenomenal. I have never seen another summary or review that pulled together so much of the relevant literature and explained it all so well. Seriously, that blog post deserves to be a review paper; it could be submitted pretty much as-is, although it would be even better with his two other Alamosaurus posts integrated (this one, and this one). It’s great work, is what I’m saying, and it needs to be acknowledged.

In particular, I was struck by the note by Anonymous in 1941 on the discovery of a cervical vertebra 1.2 meters long. I’d never heard of that ref, and I’ve never seen that vert, but at 120cm it would be in the top 7 longest cervical vertebrae on the planet (see the latest version of the list in this post), narrowly beating out the 118-cm cervical of Puertasaurus. In fairness, the preserved cervical of Puertasaurus is probably a posterior one, and more anterior cervicals might have been longer. Then again, in the big Alamosaurus neck the longest verts are pretty darned posterior, so…we need more Puertasaurus.

EDIT a few hours later: Thanks to the kind offices of Justin Tweet, I’ve now seen Anonymous (1941), and the exact wording is, “A single vertebra, or neck joint bone, is three feet across, only two inches less than four feet long, and in its present fossilized state weighs 600 pounds.” ‘Two inches less than four feet long’ is 46 inches or a hair under 117cm, which puts the supposed giant cervical just behind Puertasaurus after all, but still firmly in the top 10. And depending on how one interprets the passage in Anonymous (1941), it might not have been any bigger than BIBE 45854–see this comment for details.

Big cervical showdown. From the top left: BYU 9024, originally referred to Supersaurus but more likely representing a giant Barosaurus (137cm); the single available cervical of Puertasaurus (118cm); a world-record giraffe neck (2.4m); Alamosaurus referred cervical series BIBE 45854, longest centra are ~81cm; Sauroposeidon holotype OMNH 53062, longest centrum is 125cm. This image makes it very clear that whatever Sauroposeidon was doing, it was a way different thing from Alamosaurus.

Crucially, the longest vertebrae in the BIBE 45854 series are about 80 or 81 cm long, which means that a 1.2-meter cervical would be half again as large. That is a pretty staggering thought, and that individual of Alamosaurus–assuming it was the same taxon as BIBE 45854, and not some other, longer-necked critter–would definitely be a contender for the largest sauropod of all time.

Illustrations here are of the big Alamosaurus cervical series from Big Bend, which was comprehensively described by Ron Tykoski and Tony Fiorillo in 2016, and which we have covered in these previous posts:


  • Anonymous. 1941. Find dinosaur neck bone nearly four feet long. The Science News-Letter 39(1):6–7.
  • Tykoski, R.S. and Fiorillo, A.R. 2016. An articulated cervical series of Alamosaurus sanjuanensis Gilmore, 1922 (Dinosauria, Sauropoda) from Texas: new perspective on the relationships of North America’s last giant sauropod. Journal of Systematic Palaeontology 15(5):339-364.

Xinjiangtitan when originally described, from Wu et al. (2013)

We’re way late to this party, but better late than never I guess. Wu et al. (2013) described Xinjiangtitan shanshanesis as a new mamenchisaurid from the Middle Jurassic of China. At the time of the initial description, all of the dorsal and sacral vertebrae had been uncovered, as well as a handful of the most posterior cervicals and most anterior caudals.

Xinjiangtitan revealed, from Zhang et al. (2018)

Jump a few years forward 2018, when Zhang et al. described the complete cervical series of Xinjiangtitan, based on further excavation of the holotype (they also changed some of the element identifications in the original description). It’s pretty insane: 

  • 18 cervical vertebrae, same as Mamenchisaurus youngi, and one less than M. hochuanensis, all discovered in articulation;
  • 10 of those vertebrae have centrum lengths of 1 meter or more;
  • the longest centrum, that of C12, is 123cm long;
  • the total lengths of the separate cervical vertebrae (not articulated) add up to about 15 meters;
  • even assuming that the condyles of the vertebrae were fully buried in the cotyles, the total length of articulated neck would still be 13.36 meters. 

Now, some caveating. Zhang et al. (2018) report two different lengths for most the cervicals: a maximum centrum length, which includes the anterior condyle, and a “minimum centrum length” without the anterior condyle. Reporting cervical lengths minus the condyle is fairly common–Janensch did it for what is now Giraffatitan (“ohne condylus”), McIntosh (2005) did it for the AMNH Barosaurus, Tschopp and Mateus (2017) did it for Galeamopus pabsti, and so on. In the freely available but as-yet-not-formally-published 4th chapter of my dissertation (Wedel 2007), I referred to the length without the condyle as the “functional length”, and I explicitly assumed that it was “the length that each vertebra contributes to the total neck length”. At the time I assumed that condyles were always fully buried in cotyles in life, because I didn’t know about camel necks (see Taylor and Wedel 2013b: fig. 21 and this post). 

Why am I bringing up all these minutiae? Because I’m really interested in the actual length of the neck of Xinjiangtitan in life, and that’s not so very straightforward to figure out. I’ll start with what Zhang et al. wrote, then proceed to their measurements, and then discuss their map.

At the start of the Description section, Zhang et al. (2018: p. 3) wrote:

In SSV12001, the cervical series is almost completely articulated and is exposed laterally (Figure 2). The long neck (at least 14.9 m) is well-preserved with a total of 18 cervical vertebrae. This measurement was estimated based on the maximum centrum length including the anterior condyles with the space for the cartilage assumed.

How much space is assumed for the cartilage? They don’t say, and it’s not clear, but one reading is that they just added up the total lengths of all the cervical centra and assumed that the cotyles were completely full of cartilage. Which is not so crazy as it might sound, since that’s exactly what happens in camels. But let’s see what their tables of measurements say.

Xinjiangtitan cervical vertebra measurements, from Zhang et al. (2018)

Table 1 gives the measurements of the atlas and axis, and Table 2 gives the measurements of all the remaining cervicals. Only “minimum centrum length”–without the condyle–is reported for cervicals 4 and 5, because C3-C5 were articulated as a unit, they haven’t been separated, and without CT scanning or further prep it’s going to be impossible to determine how long they were with the condyles. However, we can infer that the condyles of C4 and C5 are buried in the cotyles of C3 and C4 because (a) only the without-condyle lengths are reported, and (b) the condyles aren’t visible in the figures. File that away, it’s going to be important.

Adding up all of the max centrum lengths, including 165mm for the axis and 30mm for the atlas, per Table 1, I get a total of 14985mm, or 14.985 meters. Because Zhang et al. were so assiduous about their reporting–they really did Measure Their Damn Dinosaur–we can estimate pretty closely how much longer that total would be if it included the condyles of C4 and C5. Subtracting the min length from the max length, we find that the condyle is 70mm long in both C3 and C6, so it’s reasonable to assume the same for the vertebrae in the middle. Adding 140mm to the earlier total gets us up to 15125mm, or 15.125 meters. That’s assuming condyles end even with the rims of the cotyles, and cotyles are completely full of cartilage.

Xinjiangtitan cervicals, from Zhang et al. (2018: fig 3)

Adding up the all of the minimum centrum lengths, again including the axis and atlas, yields a total of 13360mm, or 13.36 meters. I think this smaller total is much more likely to be the actual length of the neck in life, for three reasons:

  1. As mentioned above, the condyles of C4 and C5 of this very specimen are actually buried in the cotyles of the preceding vertebrae. So we don’t need to add any space for cartilage to the summed minimum (without condyle) lengths–there certainly was cartilage between the surfaces of the condyles and cotyles, because that’s how intervertebral joints work, but there was not enough to push the condyles back outside the cotyles, unless we want to engage in some special pleading that C3-C5 were unnaturally smooshed together.
  2. Camels notwithstanding, having the condyles buried in the cotyles is pretty standard for articulated necks of big, long-necked sauropods. In the holotype specimens of Mamenchisaurus hochuanensis and Sauroposeidon, the condyles are not visible in lateral view, because they are completely buried in the cotyles of the preceding vertebrae–see the photos in this post and on this page to confirm that for yourself. In Giraffatitan, just the edges of the condyles are visible sticking out the backs of the cotyles in some of the posterior cervicals–see this post.
  3. The 13.36-meter neck is more consistent with the map of the specimen in the ground than either the 14.9-meter or 15.1-meter totals.

A little unpacking on that last point. Using the dorsal lengths from Wu et al. (2013: table 1)–and assuming that Zhang et al. are correct, and the D1 of Wu et al. is actually cervical 18, but D11 of Wu et al. is actually D10 and D11 together, so there are still 12 dorsals–I get a total length for the articulated dorsal column of 3355mm. Dividing 13360 by 3355 yields a cervical/dorsal ratio of 3.98. Using the screenshot of the map from Zhang et al. (2018: fig. 2), I measured 1505 pixels for the summed cervicals, 380 pixels for the summed dorsals, and 112 pixels for the scale bar. Assuming the scale bar is supposed to be 1 meter (and not 20 meters or 2.0 meters as it is labeled) yields a summed cervical length of 13.4 meters, a summed dorsal length of 3.39 meters, and a cervical/dorsal ratio of 3.96–all admirably close, off by no more than 4cm across 16+ meters, if the neck in the ground was articulated condyle-inside-cotyle. If we assume the map shows a 14.9-meter neck, then both the dorsal series and the scale bar are off by about 12%, which is unreasonable given the high precision of the map if the articulated neck corresponds to the summed minimum lengths.

Mounted skeleton of Omeisaurus tianfuensis: N E C C

Bonus observation #1: the holotype of Mamenchisaurus hochuanensis has a cervical/dorsal ratio of 3.52, but in Omeisaurus tianfuensis the same ratio is 4.09. So Xinjiangtitan is actually a little shorter-necked than Omeisaurus, at least compared to the length of the dorsal series.

Bonus observation #2: the 123-cm cervical of Xinjiangtitan is only the fifth-longest vertebra of anything to date:

  1. BYU 9024, possibly referable to Supersaurus or Barosaurus: 137cm
  2. Price River 2 titanosauriform: 129cm
  3. OMNH 53062, Sauroposeidon holotype: 125cm
  4. KLR1508-77-2, Ruyangosaurus giganteus referred specimen: 124cm
  5. SSV12001, Xinjiangtitan shanshanesis holotype: 123cm
  6. MPEF-PV 3400/3, Patagotitan holotype: 120cm (+?)
  7. MPM 10002, Puertasaurus holotype: 118cm

Getting pretty crowded there in the 120s, but then a big jump to BYU 9024. I’ll have more to say on that in a second.

Just to put a bow on this section, I’m pretty confident, based on all available measurements, taphonomic evidence, and the consilience between the measurements and the map, that the holotype individual of Xinjiantitan had a neck 13.36 meters (43 feet, 10 inches) long in life. 

That’s stunning.

By comparison, the second- and third-longest complete cervical series (of anything, ever, to date) belong to Mamenchisaurus hochuanensis, at 9.5 meters (Young and Zhao 1972, and confirmed by Mike in a basement in Slovenia), and Giraffatitan at 8.5 meters for MB.R.2181 (the larger XV2 specimen would have had a 9.6-meter neck).

Some other contenders, from Taylor and Wedel 2013a (fig 3)

There were things with longer necks, for sure, but none of those necks are complete (yet). Mamenchisaurus sinocanadorum is estimated to have had a neck about 12 meters long, based on the partial cervical series of the holotype. I know there are skeletal reconstructions out there with longer necks, and I will believe them as soon as the specimens they are based on are published. In the aforementioned dissertation chapter, I estimated 11.5 meters for the neck of Sauroposeidon, assuming a brachiosaurid-like cervical count of 13. Note that Mannion et al. (2013) recovered Sauroposeidon as a somphospondyl, and a cervical count of 15 or more as a synapomorphy of Somphospondyli. Adding a couple more 1.2-meter mid-cervicals would bring Sauroposeidon up to perhaps 14 meters. The longest cervicals of Patagotitan are in about the same size class, and we don’t know the cervical count in that monster, either.

BYU 9024, with the mounted (cast, composite) skeleton of Brachiosaurus altithorax and one Mike Taylor for scale

And of course, lurking out there in crazy neck-space is BYU 9024, the immense cervical originally referred to Supersaurus, but which more likely belongs to Barosaurus, and an ungodly huge one. That critter might–might–have had a 17-meter neck.

And I gotta say, in light of Xinjiangtitan, that no longer seems so unreasonable. Because Xinjiangtitan was a big sauropod but not a monster. The dorsal length of 3.3 meters and the femur length of 1.65 meters put it in roughly the same size category as the bigger individual of Jobaria (DL 3.2m, FL 1.8m) or the AMNH 5761 Camarasaurus supremus (DL 2.5m, FL 1.8m). Let’s imagine a Xinjiangtitan with a 2.4-meter femur, the size of Patagotitan or Argentinosaurus. Assuming isometric scaling, that individual would have a 2.4/1.65 = 1.45 x 13.36 = 19.4-meter neck. 

Do we really think such animals never existed?

Food for thought: the holotype individual of Xinjiangtitan was small enough to be buried as a complete skeleton. What about the individuals that were too big to bury in one shot?

Utterly unsurprising, but still nice to see: the highly pneumatic internal structure of the vertebrae of Xinjiangtitan, from Wu et al. (2013)