New paper: pneumatic diverticula and blood vessels in the neural canals of the toothed birds Ichthyornis and Janavis
October 11, 2025
New paper out this week, open access like usual, go get it for free:
If I recall the sequence correctly, Jessie Atterholt met Dan Field at one of the recent Society of Avian Paleontology and Evolution (SAPE) meetings. Between them they spun up the idea of looking for evidence of paramedullary diverticula (PMDs) in the neural canals of some fossil birds that Dan and his collaborators and students had been studying, namely Ichthyornis and Janavis, both toothy ichthyornithines from the Late Cretaceous. This was not long after Jessie and I had our paper on PMDs in extant birds published (Atterholt and Wedel 2022), and we were interested in chasing PMDs down the tree. At the same time, Dan and his former student, Juan Benito, had a big war chest of CT scans of Ichthyornis and Janavis. So the actual work for this project was very similar to the work for Atterholt and Wedel (2022): lots of hours in front of a computer, flipping through stacks of CT slices. But I’m getting ahead of myself.
Ichthyornis you know, it’s one of the toothed birds that O.C. Marsh described in the 1870s, after basically buying the specimen out from under E.D. Cope, one of the many inciting incidents of the Bone Wars. For most of my career I simply could not keep Ichthyornis and Hesperornis straight. It has always been perversely confusing to me that the flightless swimming bird is not named “fish bird”, and the gull-like flying bird is not named for Hesperus, or Venus, a thing actually up in the sky. The “fish bird” was the flyer and the “Venus bird” was the flightless swimmer. It’s just plain backwards. (Before anyone pushes their glasses up their nose in the comments, yes, I know that Hesperornis is intended as “western bird”. Both taxa are from the West. Still confusing.)

The much larger Janavis (right) compared to the more-completely-known Ichthyornis (left). From Benito et al. (2022: fig. 1).
Janavis I was not familiar with prior to this project. It’s the sister taxon of Ichthyornis, only named in 2022 by Benito et al. Janavis was big, too, with an estimated wingspan of 5 feet, about the same as the largest extant gulls (or for me, an Oklahoma farmboy, a really big hawk). The vertebrae of Janavis are cuh-ray-zee pneumatic, totally honeycombed inside and fairly Swiss-cheesy in places on the outside, edging up to the frankly unbelievable anatomy of pelicans. Or shoebill storks, about which more in a sec.

Jessie Atterholt, Grace Burton, and me at the LACM in August, 2024. Sorry about the unfortunate non-sauropods in the background.
Grace Burton, one of Dan’s current PhD students, came over to SoCal last year to do some research at the LACM and work with Jessie and me on the IchyJan project (it only took me about half a dozen emails to realize that I was too lazy to type “Ichthyornis and Janavis” the thousand or so times I’d need to). The three of us had an enjoyable visit to the LACM Ornithology collection to find comparative specimens, some of which we ended up figuring in the new paper. And Jessie and Grace spend a LOT of time looking through CT scans. I got in on some of that, but really, Jessie and Grace did almost all the heavy lifting with both the research and the writing, so it’s only just that they’re the first two authors. This was mostly an Atterholt joint from the get-go anyway. If my interest in weird neural canal anatomy is a roaring bonfire, Jessie’s is more like the Sun.

One of the cervical vertebrae of the shoebill stork, Balaeniceps rex, LACM 116167. Check out the “bone foam” of pneumatic foramina inside the cervical rib loop and on the side of the centrum.
Of the new coauthors I picked up on this project, one is close to home: Elle Fricano, who works alongside Jessie and me as one of the anatomy faculty at WesternU. We ended up needing to scan some specimens at WesternU with our microCT machine, and Elle did virtually all of the scanning and interp, so we brought her on as an author. Elle’s own research is mostly on the evolution of the cranial base and ear region in humans and other primates, but she’s gotten into pneumaticity with a very nice paper on the human maxillary sinus (Fricano et al. 2025). She also works as a forensic anthropologist, and earlier this year she passed her forensic board exams to became the 176th Diplomate of the American Board of Forensic Anthropology — the 176th ever (full list here) — and one of only 124 active board-certified forensic anthropologists in the world. That is a heck of an achievement for anyone, but especially for someone on the tenure track, with a heavy teaching load, research, committee service, and a family. Am I bragging on my colleague? Heck yes. When a fire burns down a neighborhood out here, Elle is one of the people who goes and sifts bone shards out of the ashes and does her best to give the survivors some closure (not to mention helping investigate other deaths, ones that Nature had less of a hand in). That work is not without its costs, and I’m a little in awe of anyone who chooses to do it.

Hypothesized reconstructions of respiratory, vascular, and neurological structures in the neural canals of Ichthyornis dispar and Janavis finalidens. (a) Ichthyornis (KUVP 25472) cervical 11 showing likely arrangement of paramedullary diverticula (green) and paired extradural ventral spinal vessels (pink) relative to the spinal cord (yellow). (b) Janavis (NHMM RD 271) indeterminate mid-thoracic vertebra 1 showing likely arrangement of the extradural dorsal spinal vein (blue) relative to the spinal cord (yellow). Atterholt et al. (2025: figure 5).
Anyway: neural canals in fossil birds. We were hunting for hard evidence of pneumatic diverticula inside the neural canal, ideally unambiguous foramina opening into clearly pneumatic spaces in the neural arch or centrum. We found those foramina, and lots of other weird stuff besides. Some of the vertebrae of Ichthyornis and Janavis have bilobed neural canals, and from comparisons with extant birds we’re pretty sure the upper lobe held a big venous sinus. Crocs have one, too, in their bilobed neural canals. Most of the critters that fall evolutionarily between crocs and birds don’t have bilobed neural canals, but they may still have had big venous sinuses that simply failed to leave diagnostic traces — the curse of pneumaticity researchers extended to blood vessels.
Some of our CT scans of extant birds show that upper lobe being shared by both a big venous sinus and pneumatic diverticula, and the upper lobe is sometimes expanded into what Jessie and I nicknamed the “pneumatic attic”: a large space of variable geometry that very often has big pneumatic foramina opening into the transverse processes, postzygapophyseal rami, or neural spines. You can see the “pneumatic attic” with the pneumatic diverticula restored in a vertebra of Ichthyornis in Figure 5, above. Virtually everything we found in Ichthyornis and Janavis could be lined up 1-for-1 with an identical geometry or topology in one or another extant bird, which made us feel better about our interpretations.

Paired ventrolateral channels in Ichthyornis dispar, and examples of similar structures in extant avians. (a) Ichthyornis (ALMNH 3316) axis; note that the channel on the right has just given rise to a neurovascular foramen. (b) Ichthyornis (KUVP 25472) vertebra 11. (c) King penguin (Aptenodytes patagonicus, LACM 99854) thoracic vertebra. (d) Ichthyornis (ALMNH 3316) sacral vertebra. (e) Blue petrel (Halobaena caerulea) sacral vertebra. (f) Ichthyornis (KUVP 25472) indeterminate caudal vertebra 1. (g) Ichthyornis (KUVP 25472) indeterminate caudal vertebra 2. (h) Common loon (Gavia immer, LACM 112761) caudal vertebra. (i) Antarctic prion (Pachyptila desolata) caudal vertebra. Atterholt et al. (2025: figure 4).
One thing that needs more work is the frequent occurrence of small, paired troughs at the ventrolateral corners of the neural canal, not only in Ichthyornis and Janavis but in many extant birds as well. These troughs often bud off little vascular foramina that we can trace down into the centrum, so we’re pretty sure the troughs held blood vessels in life. A lot of vertebrates have a ladder-like arrangement of arteries in their neural canals, which could be the source of these troughs, but they might also have been produced by little basivertebral veins, which birds otherwise seem to lack. Why don’t we we just inject some dead birds, dissect them, and find out, you maybe wondering. Well, we’re gonna, at some point, but that’s at least another whole paper’s worth of work, and possibly several. We’d rather just go look up the answer, but as far as we and our reviewers could tell, no-one has ever written about these troughs and their contents before (if you know otherwise, please sing out in the comments!).
So once again, Jessie and I find ourselves needing to do novel anatomical research on living animals, partly because it’s worth doing in its own right, but also so that we can make progress on the paleontological questions that got us into this in the first place. It’s awfully hard to make informed paleobiological inferences when so much basic anatomy remains to be documented for the first time, even in extant critters. As I keep saying, a lot of this is work that anyone with sufficient time and curiosity could do, much of it inexpensively. So if you find this stuff intriguing, we’d love to have more explorers out here where the pneumatosphere intrudes into the neural-canal-iverse.

I was up inside the Utah Field House Diplodocus three weeks ago, logging pneumatic structures that no-one had documented in 125 years. More on that another time. Many thanks to John Foster for the ladder and the permission.
As for Jessie and me, this is our fifth neural-canal-related paper (see the evolving list here). We keep kicking them out the rate of one per year, which is nice and sustainable and unlikely to stop anytime soon. According to my to-do list, she and I have at least another 15 collaborative papers planned. Not all of them are about neural canals, but still… I reckon we’d better get to it.
REFERENCES
- Atterholt, Jessie, and Wedel, Mathew J. 2022. A computed tomography-based survey of paramedullary diverticula in extant Aves. The Anatomical Record 306(1): 29-50. https://doi.org/10.1002/ar.24923
- Atterholt, Jessie; Burton, M. Grace; Wedel, Mathew J.; Benito, Juan; Fricano, Ellen; and Field, Daniel J. 2025. Osteological correlates of the respiratory and vascular systems in the neural canals of Mesozoic ornithurines Ichthyornis and Janavis. The Anatomical Record. http://doi.org/10.1002/ar.70070.
- Benito, Juan; Kuo, Pei-Chen; Widrig, Klara E.; Jagt, John W. M.; Field, Daniel J. 2022. Cretaceous ornithurine supports a neognathous crown bird ancestor. Nature. 612 (7938): 100–105. doi:10.1038/s41586-022-05445-y.
- Fricano, Ellen E.I; Nguyen, Joseph; Hallal, Ryan; and Llera Martín, Catherine J. 2025. Under the surface: Correlates with maxillary sinus shape. Journal of Anatomy. https://doi.org/10.1111/joa.14283
- Marsh, O.C. 1880. Odontornithes: a monograph on the extinct toothed birds of North America. United States Geological Exploration of the 40th Parallel. Washington, DC: U.S. Government Printing Office, 201 pp.
My Constant Reader, and staying close to the work
January 16, 2025

A middle caudal vertebra of a diplodocid, presumably Tornieria africana, on display at the Museum fur Naturkunde Berlin, in left lateral view.
Quick backstory: this post at Adam Mastroianni’s Experimental History led me to this post at Nothing Human, and poking around there led me to another good’un: “Shallow feedback hollows you out”. That post really hit for me, and it made me think about SV-POW! Especially this bit:
Suppose you don’t want to lose your ability to think new thoughts and see new things. What are your options?
The best remedy is to write to the single smartest person you know who cares a lot about your topic of interest.
I have two thoughts about this. The first, which dovetails nicely with the thesis of that post, is that SV-POW! staying relatively small is probably a good thing. We’ve never written with the goal of growing our readership, and I think that’s kept us from being tempted by a lot of bad habits whose deleterious effects you can see play out over and over again across the whole internet. Our habit of posting on a completely irregular schedule on whatever topics we like has been doubly beneficial: it’s kept us sane (for reasons explored in this post), and it’s probably kept our readership low,* which has kept the temptation to write for marginal readers from ever getting off the ground. In case that sounds insulting or dismissive to our readers, let me clarify: we love our readers, and we’d rather have our little community of dedicated weirdos than any other set.
(Don’t get me wrong, I like it when one of our posts goes viral, but I like it in the same sense that I like watching a comet: it’s a cool phenomenon that I feel is beyond my influence. I enjoy it, but it doesn’t affect how I conduct myself.)
*Having written that, I wonder now if our irregular posting schedule has possibly deepened the dedication of those readers who can tolerate it — it could be a form of intermittent reinforcement, which has been implicated in gambling addiction.
That leads to my second thought: at any given time in the 17-year history of this blog, we’ve had a small but dedicated cadre of commenters, but the makeup of that group has changed over time. This has also had a salutary effect: for every post I’ve ever written here, I could be pretty sure that at least some of the regulars would see it and comment, but the one thing of which I could be absolutely certain is that the post would be seen and read by Mike. For most posts, Mike probably cares as much or more about what I’m writing than anyone else in the world, he will absolutely call me to account if he catches any weaknesses of evidence or reasoning, and he’ll do it publicly, in our own comment section. These are all good things! As my Constant Reader, Mike’s helped enforce the good habits of mind and of writing that are the subject of that Nothing Human “Shallow feedback” post.

The same Tornieria vertebra in dorsolateral oblique view, showing some pneumatic features on the lateral aspect of the neural spine. The pocks on the centrum are also raising my pneumaticity antennae, but I can’t be sure from my limited set of 16-year-old photos. When Diplodocus caudals have pneumatic features this far back in the tail, they’re more commonly on the centrum than the arch, but diverticula gonna diverticulate.
Speaking of, I also really liked this bit from the first comment on that post, by Mo Nastri:
…the details change but the general pattern is the same. In each case the [once great] intellectual in question is years removed from not just the insights that delivered fame, but *the activities that delivered insight*.
To the extent that this blog has escaped enshittification, it’s probably because Mike and I are not removed from the activities that deliver insight. We care more about sauropod vertebrae (and pig skulls, etc.) than we do about clicks. And at this point, I’m confident that we always will. If we were ever in danger of click-maximizing behavior, it was probably back in the early days, and even then the risk was minimal. We love our weird little niche blog just as it is, weird and niche-y and little.
The possibly-surprising conclusion I’m building toward is that we’ve probably made SV-POW! a better experience for our readers (minimally, in that it still exists to be read) by not caring about our readership, and by not writing to please or impress anyone other than ourselves and each other. And that in turn has kept SV-POW! viable for us as well.
So if you’re here, great! We’re happy to have you — as an interested person, rather than a click. If you like what we’re doing, stay tuned. We’re gonna do a lot more of the same.
To what extent is science a strong-link problem?
October 30, 2024
Here’s a fascinating and worrying news story in Science: a top US researcher apparently falsified a lot of images (at least) in papers that helped get experimental drugs on the market — papers that were published in top journals for years, and whose problems have only recently become apparent because of amateur sleuthing through PubPeer.
I’m going to wane philosophical for a minute. In general I’m very sympathetic to Adam Mastroianni’s line “don’t worry about the flood of crap that will result if we let everyone publish, publishing is already a flood of crap, but science is a strong-link problem so the good stuff rises to the top”. I certainly don’t think we need stronger pre-publication review or any more barrier guardians (although I have reluctantly concluded that having some is useful). But when fraudulent stuff like this does in fact rise to the top in what seems to be a strong-link network — lots of NIH-funded labs, papers in top journals (or, apparently, “top” journals) — then I despair a bit. Science has gotten so specialized that almost anyone could invent facts or data within their subfield that might pass muster even with close colleagues (even if those colleagues aren’t on the take, he said cynically — there is a mind-boggling amount of money floating around in the drug-development world).
Immediate thought experiment: could Mike or I come up with material for a blog post or paper that would be false but good enough to fool the other? Given how often we find surprising or even counterintuitive results, I think possibly so. I’m not particularly motivated to run the experiment when we’re already digging out from a deep backlog of started-but-never-finished papers, but it remains a morbidly fascinating possibility.
Anyway, one problem is that “top” journals have a lot of fraudulent or at least incorrect science in them, roughly corresponding to their impact factors. Now, you might say “yeah but the positive correlation means bad actors get caught”, to which I’d reply “not fast enough” and “how do you know we’re catching all of them?”
Sinking feeling
There’s another problem, I don’t know if it’s equal-and-opposite but it definitely exists: good science that doesn’t float to the top. Here are a couple of quick examples from my neck of the woods:
Working from very little evidence by modern standards, Longman (1933) had correctly figured out that pneumatic sauropod vertebrae come in two flavors, those with a few large chambers and those with many small chambers. He called them “phanerocamerate” and “cryptocamerillan”, corresponding to the independently-derived modern terms “camerate” for the open-chambered form and “camellate” or “somphospondylous” for the honeycombed one. As far as I have been able to determine, nobody paid any attention to this before Wedel (2003b) — Longman’s work on vertebral internal structure wasn’t mentioned or cited by Janensch in the 1940s or Britt or anyone else in the 1990s. To be clear, I’m not putting myself forward as a better researcher than anyone that came before. I just got lucky, to have read a fairly obscure paper while I had my antennae out for any possible mention of pneumaticity.
Speaking of Janensch, his 1947 paper on pneumaticity in dinosaurs was pretty much ignored until the 1990s and early 2000s.

OMNH 1094, a cervical centrum of an apatosaurine, and a crucial player in the Wedel origin story — this was the first vertebra of anything other than Sauroposeidon that Kent Sanders and I scanned.
I owe my career to the Dinosaur Renaissance
Here’s what bothers me about this: I made my career studying pneumaticity in sauropods, buoyed in large part by the fact that I stumbled backwards into a situation where I had access to a big collection of sauropod bones (at the OMNH), free time on a CT scanner (at the university hospital), and a curious and collaborative radiologist (Kent Sanders). But you don’t need a CT scanner to study pneumaticity, as John Fronimos has convincingly demonstrated (see Fronimos 2023 and this post). So why didn’t the revolution in sauropod pneumaticity happen in 1933 or 1947? Or, heck, in 1880 — Seeley and Cope and Marsh and many others recognized that sauropods had highly chambered vertebrae.
I think the most likely explanation is that at the time no-one cared. Pneumatic vertebrae in sauropods were possibly interesting trivia, but sauropods were an evolutionary dead end and so their vertebrae couldn’t tell us anything important about evolutionary success. These attitudes may not have been universal, but they were certainly prevailing.
I had the good fortune to come along at a time when there was renewed interest in dinosaur paleobiology, particularly any characters or body systems shared between non-avian dinosaurs and birds. Suddenly pneumaticity wasn’t some obscure bit of trivia, but the skeletal footprint of a bird-like respiratory system that was potentially a key adaptation for sauropods (Sander et al. 2011) and possibly for dinosaurs more generally (Schachner et al. 2009, 2011). And dinosaurs weren’t any more of an evolutionary dead end than we are, they just happened to mostly not fit into small holes or deep water when the asteroid hit. (Let’s heat the atmosphere to 400F for a few hours and then make the world dark for a few months or years and then we can talk about evolutionary dead ends.) So adaptations that facilitated dinosaurosity might tell us something about evolutionary success after all.
What are you doing in that cell?
Having a successful career because I happened to hitch a ride on a wave of renewed interest in dinosaur paleobiology is certainly nice, but also worrisome. If it takes 70 or 100 years for the good science to float to the top, does that really count? Whatever convection cells push the good science toward the top would ideally work more like a cook pot on a rapid boil, and not like the imperceptible roiling of Earth’s mantle. So ask yourself: what’s still on its way up to the top right now, that no-one has clocked yet? What’s the Longman (1933) of 2024 — the seemingly incidental observation that is going to seem prophetic in a few decades? Or worse, what was the Longman (1933) of 1994 or 2004, the solid paper that attracted no attention and won’t for another half century?
The convection cell metaphor is particularly apt because a lot of science is siloed. A good idea — say, that the peroneus tertius muscle occurs at a lower frequency in monkeys and apes than in humans, and this tells us something about its evolution — may rise to the top in one cell (comparative anatomy), but not make it over to the neighboring cell (clinical anatomy), where all the happy little molecules think that peroneus tertius is a muscle unique to humans (if you have no idea what I’m on about, see the second numbered point in this post).
So if you want to do good work — in this metaphor, to be at the top where the good science floats (eventually, alongside a seasoning of not-yet-unmasked bad science) — then I think you have to be aware that other cells exist, and occasionally peer into them, if for no other reason than to make sure you don’t accept an idea that’s already been debunked over there. And you need to read broadly and deeply in your own cell — there’s almost certainly valuable stuff you don’t know because the relevant works are stuck to the bottom of the pot. Go knock ’em loose.
References
- Fang, F.C. and Casadevall, A. 2011. Retracted science and the retraction index. Infection and Immunity 79(10):3855-3859.
- Fronimos, John A. 2023. Patterns and function of pneumaticity in the vertebrae, ribs, and ilium of a titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous of Texas, Journal of Vertebrate Paleontology 43:2. DOI: 10.1080/02724634.2023.2259444
- Janensch, W. 1947. Pneumatizitat bei Wirbeln von Sauropoden und anderen Saurischien. Palaeontographica, Supplement 7, 3:1–25.
- Longman, H. A. 1933. A new dinosaur from the Queensland Cretaceous. Memoirs of the Queensland Museum 10:131–144.
- Sander, P.M., Christian, A., Clauss, M., Fechner, R., Gee, C.T., Griebeler, E.M., Gunga, H.C., Hummel, J., Mallison, H., Perry, S.F. and Preuschoft, H. 2011. Biology of the sauropod dinosaurs: the evolution of gigantism. Biological Reviews 86(1):117-155.
- Schachner, E.R., Lyson, T.R. and Dodson, P., 2009. Evolution of the respiratory system in nonavian theropods: evidence from rib and vertebral morphology. The Anatomical Record 292(9): 1501-1513.
- Schachner, E.R., Farmer, C.G., McDonald, A.T. and Dodson, P., 2011. Evolution of the dinosauriform respiratory apparatus: new evidence from the postcranial axial skeleton. The Anatomical Record 294(9): 1532-1547.
The pneumatic rib evolution figure in a more useful format
October 25, 2024
Here’s a funny thing I hadn’t given much thought to until recently: virtually all journals, even the born-digital variety, have pages in portrait mode for easy printing on 8.5×11 or A4 paper. And many offer a column-width option for figures. So if you want to line up a whole bunch of stuff for easy comparison, for a paper it’s usually easier to orient a figure vertically, like so:
And here it is in context on the page:
But virtually all slide presentations use a landscape format, 4:3 for a long time but often 16:9 these days to accommodate wider screens, or phones and tablets in landscape mode. For this a figure much taller than wide is usually not a good use of space, and may present at too small a scale to be readable.
I ran into this last week while prepping a presentation on my research for an anatomy department meeting at work. I wanted to use that King et al. figure because it summed up so much of the paper in one image, but the only version I had was the skyscraper version we’d used in the JVP paper. So I went into GIMP and rotated the image and every element within it by 90 degrees, to produce this landscape version:
I was presenting to an intellectually diverse audience, most of whom do not work on dinosaurs, so I added little silhouettes (my own, cribbed and hacked from all kinds of older work) to make it all more explicable:
This is all my original work, and I’m letting it out in the world here in case anyone else wants to use it. CC-BY like everything else on this blog. FWIW I think mamenchisaurs and diplodocids held their necks elevated — the baseline alert posture for extant tetrapods — I was just moving quickly and more concerned with getting little doodads for all the genera than with any paleobiological implications.
So now I’m wondering if there are any figures in old papers that I’ve avoided putting in talks, possibly subconsciously even, because they’re the wrong shape. Not that I need to do any more navel-gazing than I already do, but maybe something for me to keep an eye out for when I have reason to go back to them (which is often — they’re thought archives).
The more forward-looking takeaway is that if you have to make a taller-than-wide figure to fit a journal page, consider making a wider-than-tall version at the same time to throw into your talks — or vice versa if you’re making the talk first. It’s a time investment for sure, but it may be easier while all the bits are fresh in your head and you have all the elements in separate layers or whatever. Hopefully you already back up the uncompressed versions of all your figures, but Past Matt didn’t always do that, so at least be smarter than that guy!

Tate v2610, a sauropod dorsal rib. Check out the nice deep pneumatic fossa a little way down from the tuberculum of the rib (upper left in the photo).
Parting shot (and an excuse to post a photo for Fossil Friday): on my Tate trip this summer I hit a gang of museums, and everywhere I went I found pneumatic sauropod ribs. I think there are a lot more of these things out there than most folks have appreciated. I’m proud of my recent pneumatic rib papers (Taylor et al. 2023 and King et al. 2024), but I hope they are the just the start of something.
And because I picked that photo: you know what institution has a ton of super-interesting, well-preserved, well-prepped, not-yet-published-on sauropod vertebrae and ribs in a really nicely appointed collections room in an awesome museum run by a small team of excellent human beings? The Tate Geological Museum, that’s who. If you can get yourself to Casper and you have a legit research interest, go check out their collections, there’s SO MUCH good stuff in there. I myself will be back as soon as it can be conveniently arranged.
References
- King, J.L., McHugh, J.B., Wedel, M.J., and Curtice, B. 2024. A previously unreported form of dorsal rib pneumaticity in Apatosaurus (Dinosauria: Sauropoda) and implications for pneumatic variation among diplodocid dorsal ribs. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2024.2316665
- Taylor, Michael P., and Matthew J. Wedel. 2023. Novel pneumatic features in the ribs of the sauropod dinosaur Brachiosaurus altithorax. Acta Palaeontologica Polonica 68(4): 709–718. doi:10.4202/app.01105.2023
If I could dissect a sauropod…
September 12, 2024
Luke Horton asked in a comment on a recent post:
Given the chance to examine a titanosaur cadaver with your hypothetical army of anatomists, what would you look for first?
*FACEPALM* How we’ve gone almost 17 years without posting about a hypothetical sauropod dissection is quite beyond my capacity. I am also contractually obligated to remind you that the TV show “Inside Nature’s Giants” shows dissections of a whale, elephant, giraffe, tiger, anaconda, giant squid, etc., so it’s probably the closest we’ll ever get. Go look up photos of Dr. Joy Reidenberg standing, um, amidst a partially-dissected whale, or just watch that episode, and your sauropod-dissection-visualizer will be properly calibrated.
To get back to Luke’s question, there are loads of interesting things that could be dissected in a sauropod, but since the remit here is Matt Wedel x titanosaur, there’s only one possible answer: the lung/air sac system and its diverticula. For several reasons:

Hypothetical reconstruction of the lungs (red) and air sacs (blue, green, and gray) in Haplocanthosaurus CM 879. I’d love to know how close this is to reality. Wedel (2009: fig. 10).
First and most obviously, I’ve spent the last quarter-century trying to infer as much as possible about the respiratory systems of sauropods based on the patterns of pneumaticity in their skeletons, and I’d kill for the opportunity to check the accuracy of my inferences — and those of all my fellow-travelers in the sauropod and dinosaur respiration biz, like Daniela Schwarz and Emma Schachner and Tito Aureliano and many others.

Sauropod respiratory system modeled on that of a bird. I’ll bet the correspondence wasn’t this close. (Also, since making this figure 20 years ago, I’ve learned that the abdominal air sacs of ostriches are actually rather small, although the perirenal, femoral, and subcutaneous diverticula of the abdominal air sacs are extensive; see Bezuidenhout et al. 1999). Wedel and Cifelli (2005: fig. 14).
Second, I am intrigued/haunted by the possibility that extant birds might not represent the apex of saurischian lung/air sac evolution. Birds survived the K-Pg disaster because they were small; respiratory efficiency had little or nothing to do with it (evidence: all the other small-bodied tetrapods that survived, like the many, many squamate and mammalian lineages). To me it would be a wild coincidence if the tiny dinosaurs that survived also just happened to be The Bestest (TM) at some anatomical/physiological thing unrelated to their survival. In fact, given how sensitive birds are to airborne dust and ash, I wonder if their fancy lungs weren’t more of a hindrance than a help in the dusty, sooty, iridium-laced post-impact world. Anyway, there are interesting clues that the air sac systems of extant birds are just one subset of a much greater original diversity, like most (all?) birds starting out embryologically with a dozen or so air sacs, which get simplified to the usual 9 or fewer by fusions. What did other dinosaurs do with their 12 (or more?) air sacs? If any dinosaurian clade was going to push the capabilities of the “avian” lung/air sac system in interesting directions and to fascinating extremes, sauropods seem like a good bet.

Rib articulation angles in the dorsal vertebrae of (a) Lufengosaurus, (b) Diplodocus, (c) Haplocanthosaurus, (d) Tyrannosaurus, and (e) an ostrich. Anterior is to the right. Diplodocus and Haplocanthosaurus are pretty wildly different considering they coexisted in the Morrison. I really gotta write a whole post about that. Boisvert et al. (2024: fig. 12).
So I’m intrigued by the idea that extant birds show us one way that a saurischian lung/air sac system can work, but don’t exhaust the territory, anymore than kangaroos show us all the ways that mammals can reproduce. Maybe sauropods had even better lungs than birds! Or maybe not. Likely they were doing their own weirdly specialized thing — or many weirdly specialized things — that left few to no diagnostic traces in their skeletons. We can be pretty confident that at least some of the pneumatic diverticula of sauropods worked essentially identically to how they do in birds (see Woodruff et al. 2022 and this post), and mid-dorsal pneumatic hiatuses in juvenile sauropods — predicted by me in 2003, found by Melstrom et al. (2016) and Hanik et al. (2017) — suggest that their air sac systems were broadly comparable. On the other hand, the variety of rib articulation angles just within Morrison sauropods tells us they weren’t all ventilating their air sacs in quite the same way (Boisvert et al. 2024), despite broad similarities with other dinos at the levels of rib osteology (Wang et al. 2023) and whole-thorax construction (Schachner et al. 2009, 2011). (Aside: why the hell didn’t I work a citation of Wang et al. 2023 into the Dry Mesa Haplo paper? I can only conclude that I am at least occasionally an idiot.) Whatever was going on, I’m pretty sure sauropods didn’t look exactly like 60-ton turkeys on the inside, but we don’t have a ton of real data on how they differed. It would be amazing to find out.

The mounted Rapetosaurus skeleton at the Field Museum, traced from a photo. Specific weird things to note: neck about twice as long as tail, cervical vertebrae about twice as tall as dorsals, and smallish pelvic bones relative to hindlimbs (= skinny posterior abdomen, at least dorsoventrally). See this post for details.
Third, if any sauropods were going to rival or exceed birds in fancy under-the-hood anatomical and physiological adaptations, my money would be on titanosaurs. They were morphologically disparate, phylogenetically diverse, geographically widespread, they independently evolved to giant size more times than any other sauropod clade, and their growth rates were wild. I’d dissect any sauropod I got access to (uh duh), but a titanosaur would be particularly appealing. Which titanosaur? Probably Rapetosaurus: we know it grew very fast early on (Curry Rogers et al. 2016, and see implications for the nervous system in Smith et al. 2022), it had a highly pneumatic vertebral column (O’Connor 2006), its body proportions were pretty wacky, and it had other features of interest to me, like expanded neurocentral joints (see Wedel and Atterholt 2023 and this post) and neural canal ridges (see Atterholt et al. 2024 and this post).

I used this photo of a Rapetosaurus caudal vertebra a few posts ago to illustrate the neural canal ridges, but — like many other sauropods — it also has very expanded neurocentral joints forming boutons. From Curry Rogers (2009: fig. 27).
Oh, and if I got to dissect more than one sauropod, the rest of my top 5 choices in order would be:
- the owner of BYU 9024 (Supersaurus? Giant ancient individual of Barosaurus? Are those even different things? Dissecting this critter could tell us!), Barosaurus being the most diplodocid-y and least titanosaur-y neosauropod I know of, and BYU 9024 being from a hellaciously big individual no matter what its classification;
- the Snowmass Haplocanthosaurus, because I have just so many questions about all the weird stuff going on with its tail (see Wedel et al. 2021 and this post for starters);
- Omeisaurus or Xinjiangtitan, to represent a maximally derived-but-also-weird non-neosauropod;
- Sauroposeidon, for obvious emotional reasons (but not enough to dethrone the others).
After that? Probably Isanosaurus or Melanorosaurus or something else waaaay down the tree, so I could see how much of the sauropod kit was in place from the get-go (probably most of it).

Bone vs joint space in the proximal caudals of the Snowmass Haplocanthosaurus. I’d give one non-essential organ to dissect that tail!
And after the respiratory system, next up for me would be the spinal cord and any related morphological specializations of the neural canal — see Table 3 in Atterholt et al. (2024) for a running tally, and this page. Then intervertebral joints, digestive tract, and reproductive system (neither of the last two leave anything useful in the way of skeletal traces), in that order. Arguably the intervertebral joints would be a bigger score for sauropod paleobiology than spinal cord stuff, but maybe not, and having squelched my emotional pick among sauropod taxa, I’m letting my emotions rule when choosing body systems to dissect. I also am intensely interested in the possibility of protofeathers in sauropods, but you don’t have to dissect those, you can just see if any are present, so I’d cheat a little and note any integumentary specializations en passant. (Remember than an animal can have hairs without being hairy [naked mole rats, rhinos, manatees, dolphins], ditto for feathers.)
So that’s the sauropod and the body system I’d dissect first, if given the chance. What’s your answer?
References
- Atterholt, J., Wedel, M.J., Tykoski, R., Fiorillo, A.R., Holwerda, F., Nalley, T.K., Lepore, T., and Yasmer, J. 2024. Neural canal ridges: a novel osteological correlate of postcranial neuroanatomy in dinosaurs. The Anatomical Record, 1-20. https://doi.org/10.1002/ar.25558
- Bezuidenhout, A.J., H.B. Groenewald, and J.T. Soley. 1999. An anatomical study of the respiratory air sacs in ostriches. Onderstepoort Journal of Veterinary Research 66:317-325.
- Boisvert, Colin, Curtice, Brian, Wedel, Mathew, & Wilhite, Ray. 2024. Description of a new specimen of Haplocanthosaurus from the Dry Mesa Dinosaur Quarry. The Anatomical Record, 1–19. http://doi.org/10.1002/ar.25520
- Curry Rogers, Kristina. 2009. The postcranial osteology of Rapetosaurus krausei (Sauropoda: Titanosauria) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology 29:1046-1086.
- Curry Rogers, K., M. Whitney, M. D. D’Emic, and B. Bagley. 2016. Precocity in a tiny titanosaur from the Late Cretaceous of Madagascar. Science 352:450–454.
- 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.
- 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
- O’Connor, P.M. 2006. Postcranial pneumaticity: an evaluation of soft-tissue influences on the postcranial skeleton and the reconstruction of pulmonary anatomy in archosaurs. Journal of Morphology 267: 1199-1226.
- Schachner, E.R., Lyson, T.R. and Dodson, P., 2009. Evolution of the respiratory system in nonavian theropods: evidence from rib and vertebral morphology. The Anatomical Record 292(9): 1501-1513.
- Schachner, E.R., Farmer, C.G., McDonald, A.T. and Dodson, P., 2011. Evolution of the dinosauriform respiratory apparatus: new evidence from the postcranial axial skeleton. The Anatomical Record 294(9): 1532-1547.
- Smith, Douglas H., Rodgers, Jeffrey M., Dollé, Jean-Pierre, and Wedel, Mathew J. 2022. Giraffes vs. blue whales vs. dinosaurs: contest reveals which one builds its nervous system fastest to evade predators. Scientific American, https://www.scientificamerican.com/article/giraffes-vs-blue-whales-vs-dinosaurs-contest-reveals-which-one-builds-its-nervous-system-fastest-to-evade-predators/
- Wang, Y.Y., Claessens, L.P. and Sullivan, C., 2023. Deep reptilian evolutionary roots of a major avian respiratory adaptation. Communications Biology, 6(1), p.3.
- Wedel, M.J. 2003a. Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs. Paleobiology 29:243-255.
- Wedel, M.J. 2009. Evidence for bird-like air sacs in saurischian dinosaurs. Journal of Experimental Zoology 311A:611-628.
- Wedel, M.J., and Atterholt, J. 2023. Expanded neurocentral joints in the vertebrae of sauropod dinosaurs. In Hunt-Foster, R.K., Kirkland, J.I., and Loewen, M.A. (eds), 14th Symposium on Mesozoic Terrestrial Ecosystems and Biota. The Anatomical Record 306(S1):256-257.
- Wedel, M.J., and Cifelli, R.L. 2005. Sauroposeidon: Oklahoma’s native giant. Oklahoma Geology Notes 65 (2):40-57.
- 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.
- Woodruff, D. Cary, Wolff, Ewan D.S., Wedel, Mathew J., Dennison, Sophie, and Witmer, Lawrence M. 2022. The first occurrence of an avian-style respiratory infection in a non-avian dinosaur. Scientific Reports 12, 1954. https://doi.org/10.1038/s41598-022-05761-3
All the SV-POW! videos, and other stuff on the sidebar
June 25, 2024

BYU 11505, a caudal vertebra of a diplodocid from Dry Mesa, in posteroventral view. Note the paired pneumatic foramina on the ventral surface of the centrum.
If you want to find the paleontology and anatomy videos that Mike and I have done (plus one video about open access), they have their own sidebar page now, for your convenience and for our own. It’s, uh, just to the right of where your eyes are pointing right now. You know what, I’m sure you’ve got this.
Make a better living with extensively curated sidebar pages
In fact, we’ve added a few sidebar pages in the comparatively recent past (for a blog in its 17th year). In addition to the video page, we now have some project-specific pages, namely Mike’s Supersaurus, Ultrasaurus and Dystylosaurus in the 21st Century and Mike’s open projects pages, and my own Neural canal projects page. For now the global list of Haplocanthosaurus posts lives on the page for the Wedel et al. (2021) Haplocanthosaurus neural canal paper, but I imagine it’s only a matter of time until I add a page just to track all my business with Haplocanthosaurus. Also, ugh, I still have a few papers that I’ve blogged about, but which don’t have pages, and are therefore just that much harder to find.
And of course we still have all the old standbys: Tutorials, Things To Make and Do, The Shiny Digital Future, and so on.
It might seem kinda dumb to do a post alerting people to stuff that they can find for themselves, but the whole point of having the sidebar pages is that SV-POW! has gotten to be rather unmanageably vast, and anything that helps us — or even you! — get to the right posts quickly is a welcome assist.
The photo up top has nothing to do with any of this, I just thought it would be a fun way to meet our titular mandate.
My friend and frequent collaborator (one, two, three) Tito Aureliano invited me to give a talk on his YouTube channel, I suggested pneumaticity and gigantism, and here we are. There’s a decently lengthy Q&A, moderated by Tito, after the talk itself. Hilariously — and kindly — one of the commenters pointed out that I hadn’t explicitly answered the titular question in the talk, so I took a stab at it in the Q&A. I come on about 1:40, the talk starts about 5:20, and the Q&A starts at 1:24:30.
If you’ve seen any of my pneumaticity talks since, er, I gave my dissertation finishing talk in 2007, you’ve seen at least a few slides of this. But you won’t have seen all of them before, because a good number of them didn’t exist; this is sort of a Frankenstein stitched together from previous talks, new observations, and trying to think about the future. In particular, almost my entire 2012 SVPCA talk is crammed in near the end.
If in the talk I sound less certain about some things than I have in the past, that’s accurate. In the past few years I feel like I’ve accumulated a lot of interesting pieces (most of my post-2020 papers), and I’m in quest of a new synthetic foundation for my work (e.g., Taylor and Wedel 2021, this post), but I’ve also Seen Things that have rocked my certainty about my own level of understanding (e.g., Aureliano et al. 2023, this post). I’m cool with that. I think that whatever comprehension of pneumaticity I’m questing toward is going to have to emerge inductively from all the pieces, new and old, that I and others are producing. That’s an exciting prospect, and I’m having enough fun with the individual Legos that I’m in no tearing rush to guess what the final product will look like.
Many thanks to Tito for the invitation. After having just given a big talk that was a little speculative and a little outside my wheelhouse, it was nice to come back to home base, but hopefully still give people some useful things to take away. Time will tell.

BYU 12613, a very posterior cervical (probably C14 or C15) of a diplodocine sauropod, probably Kaatedocus or Diplodocus, from Dry Mesa, original fossil and 50% scale 3D print. The real bone has a mid-height centrum length of 270mm, compared to 642mm for C14 of D. carnegii.
I intended for the next post to be a follow-up on the new paper describing the Dry Mesa Haplocanthosaurus, as I hinted/promised in the last post. But that post is still gestating, there’s a lot of other cool stuff happening right now, and I don’t want to put off posting about it and risk never getting around to it.
Pneumatic diverticula in birds on the cover of Nature
I’m probably getting to be a crank on the subject of how pneumatic diverticula in birds are so grotesquely understudied. F’rinstance: the poultry industry is a $77 billion per year concern in the US alone, and the lung/air-sac system and its diverticula are a route for potentially lethal infections (which also affected sauropods), so you’d think we’d have the diverticular system of chickens and turkeys completely mapped, and its development fully charted. But we don’t!
See? Crank!
Anyway, Emma Schachner — who’s been doing awesome work in the arena of reptile and bird respiration for years (see here, here, and the comment thread here, for starters) — and colleagues just put bird diverticula on the map in a most spectacular fashion, with a cover article in Nature.
The short, short version is that Schachner et al. surveyed the sub-pectoral diverticulum (SPD) in 68 species of birds (in 42 families and 25 orders), and found that it was present in all soaring taxa, where it evolved at least 7 times, but absent in non-soarers. Furthermore, the SPD is in the right place to improve the mechanical advantage of the pectoralis muscles, which have a different architecture in soaring taxa, one that appears to be adapted in concert with the SPD for the particular demands of soaring flight. Schachner et al. illustrate their findings with just a ton of cool dissection photos, CT slices, and 3D reconstructions, in both the paper proper and the SI. Happily, the paper is a proper publication, 6 pages long and with plenty of detail, and not cut down to a glorified abstract.
Many things make me happy about this paper: the references to Owen (1836) and Strasser (1877), who independently suggested that the diverticula of birds might positively affect their flight dynamics; a strong team of authors taking a largely neglected anatomical system and spinning it into scientific gold; and the participation of my friends Raul Diaz and Jessie Atterholt. Together with our 2022 paper in the Anatomical Record, this is Jessie’s second taxonomically broad survey of a previously under-documented diverticular system in birds in just over two years, which is a heck of a (ahem) feather in her cap.
Given that birds have a whole internal zoo of diverticula that go between their muscles, among their viscera, under their skin, and into their bones — almost all of which are known from a bare handful of documented examples — I’m sure that there are many, many more exciting discoveries to make in this space. As Schachner et al. put it, “The discovery of a mechanical role for the respiratory system in avian locomotion underscores the functional complexity and heterogeneity of this organ system, and suggests that pulmonary diverticula are likely to have other undiscovered secondary functions.”
(If you’re thinking of not working on pneumaticity because some people are already working on [their own little corners of] it, perish the thought. At the current rate it could take decades just to document where the diverticula are and what they look like, let alone their functional implications. If the world can accommodate a new theropod phylogeny every couple of weeks, it can stand a lot more work on pneumaticity in birds and other dinos.)
Video: all the Oklahoma dinosaurs
My longtime friend and mentor, Kyle Davies, is the head preparator at the Sam Noble Oklahoma Museum of Natural History in Norman, where I did my undergrad and master’s work. Kyle is a phenomenally skilled morphologist, and if he needs something for education or exhibit that he can’t otherwise get hold of, he’ll just sculpt it himself — we’ve featured his work before (here and here). He recently gave a brown-bag lunch talk reviewing all of Oklahoma’s dinosaurs, and it’s just been posted to YouTube. Go have fun.
More travel and collections pics real soon

Luke Horton helping me get a shot of the right side of the ‘Jimbo’ Supersaurus dorsal on display at the Tate Geological Museum in Casper. The left side of this cast is visible in this post.
I just got back from a crazy-awesome research trip that I structured around the Tate Geological Museum’s 2024 summer conference. I got to spend time seeing the exhibits and working in the collections of a host of institutions, including:
- the University of Wyoming Geological Museum in Laramie,
- the Tate Museum in Casper,
- the Wyoming Dinosaur Center in Thermopolis,
- the Natural History Museum of Utah in Salt Lake City,
- the Museum of Ancient Life in Lehi, Utah,
- and — chronologically last but certainly not least — the BYU Museum of Paleontology in Provo.
At BYU I got three days to roam through the collections with Colin Boisvert, Brian Curtice, Ray Wilhite, and Gunnar Bivens. It was easily one of the most productive research trips I’ve ever had, rivaled only by the 2016 Sauropocalypse with Mike. In fact, we’d hoped that Mike would get to join me for part or all of the trip, but as luck would have it he had day job trips of his own in the same time frame. He did at least get to see the mounted cast of D. carnegii in Vienna, which he was keen to see.
This trip also had this in common with the 2016 Sauropocalypse: everywhere I went, curators, collections managers, and students were unfailingly kind, hospitable, and generous with their time and knowledge. Thanks in particular to Julian Diepenbrock, Laura Vietti, and Whitney Worrell in Laramie; JP Cavigelli, Dalene Hodnett, Shaedon Kennedy, and Rachel Stevens in Casper; Tom Moncrieffe and all the staff in Thermopolis; Carrie Levitt-Bussian at the NHMU; Rick Hunter and April Hullinger in Lehi; and Rod Scheetz, Colin Boisvert, Jacob Frewin, and Isaac Wilson at BYU.
A special thanks to Luke Horton, who is currently an undergrad at Texas A&M. He made it out to Casper for the Tate conference and field trips, and he stuck around for a day afterward to assist me in collections. Given his passion for paleontology and his work ethic, I expect you’ll be hearing more about Luke in the not-too-distant future.
The upshot of all of this is that I have roughly a million cool things to post from the trip, many of which I’ll no doubt forget about or never get around to, but I will make an effort to convert trip photos into blog fuel this summer. The photo up top is the first snowball in what will hopefully become an avalanche. At BYU I was cruising down one of the aisles of sauropod vertebrae (yes, at BYU they have literally aisles of sauropod vertebrae — heaven!) and I did a double-take: it was my old friend BYU 12613! Mike and I figured that vert in our 2013 neural spine bifurcation paper, and I’d used the 50% scale 3D print in my Dolly video. I’d brought the print along on the trip as a handy visual and tactile aid for introducing people to sauropod cervical morphology, and I’d passed it around for show-and-tell during my Tate keynote talk. I couldn’t resist putting the real fossil and the 3D print together for a photo op. Here’s one more for the road, in postero-dorsal view this time:
In addition to blog posts, you’ll be seeing photos from this trip in presentations and papers as soon as it can be decently arranged. Stay tuned!
References
- Atterholt, Jessie, and Wedel, Mathew J. 2022. A computed tomography-based survey of paramedullary diverticula in extant Aves. The Anatomical Record 306(1): 29-50. https://doi.org/10.1002/ar.24923
- Schachner, Emma R., Andrew J. Moore, Aracely Martinez, Raul E. Diaz Jr, M. Scott Echols, Jessie Atterholt, Roger W. P. Kissane, Brandon P. Hedrick, & Karl T. Bates. 2024. The respiratory system influences flight mechanics in soaring birds. Nature 630: 671-676. (link)
- Wedel, M.J. 2024. The sauropod heresies: evolutionary ratchets, the taphonomic event horizon, and all the evidence we cannot see. The Jurassic: Death, Diversity, and Dinosaurs, 28th Annual Tate Conference abstract book: 35-38.
- Wedel, Mathew J., and Michael P. Taylor. 2013. Neural spine bifurcation in sauropod dinosaurs of the Morrison Formation: ontogenetic and phylogenetic implications. Palarch’s Journal of Vertebrate Palaeontology 10(1):1-34. ISSN 1567-2158.

Pneumatic dorsal ribs in a selection of ornithodiran taxa. Clades that lack pneumatic ribs have been omitted, including non-dinosaurian dinosauromorphs, ornithischians, all early diverging sauropodomorphs, and numerous sauropods. The only included clade for which dorsal rib pneumaticity might be synapomorphic is Titanosauriformes. Phylogenetic relationships of the sauropods are based on Mannion et al. (2013) for titanosauriforms (note that the position of Brontomerus is uncertain), Tschopp et al. (2015) for diplodocoids, and Zhang et al. (2022) for Xinjiangtitan. Ribs are not shown to scale. Ribs traced from Butler et al. (2009:fig. 1b, Raeticodactylus), Campana (1875:fig. 8, Gallus), Madsen Jr. and Welles (2000:plate 19, Ceratosaurus), Zhang et al. (2022:fig. 14, reversed, Xinjiangtitan), a photo of WDC-DMJ-021-134 provided by David Lovelace (Supersaurus; see Lovelace et al., 2007), Gilmore (1936:plate 29, reversed, Apatosaurus), Riggs (1904:plate 75, Brachiosaurus), Janensch (1950:fig. 108, reversed, Giraffatitan), Wilson and Upchurch (2009:fig. 21, reversed, Euhelopus), Taylor et al. (2011:fig. 7, Brontomerus), and Curry-Rogers (2009:fig. 30, Rapetosaurus). King et al. (2024:fig. 3).
New paper out today with Logan King, Julia McHugh, and Brian Curtice, on pneumatic ribs in Apatosaurus and Brontosaurus (King et al. 2024).
This one had an unusual gestation. In the summer of 2002 2022 I did a road trip to Utah and western Colorado with my friend and frequent collaborator Jessie Atterholt. We did day trips to other collections, but we used Dinosaur Journey in Fruita as home base, and spent most of our time there. That’s where I first met Logan King, who was then recently graduated from Mike Benton’s lab at Bristol. Logan was spending the summer working for Julia McHugh at the Mygatt-Moore Quarry, and Logan and Julia were writing up MWC 9617, a sauropod rib from Mygatt-Moore with interesting pneumatic features.
Now, I had been interested in pneumatic ribs in sauropods for many years, and I’d amassed a war chest of published examples. But I had to admit to myself that the hypothetical pneumatic rib paper I’d been planning was simply never going to be my top priority, and therefore I was never going to actually start it, much less finish it. Logan and I hit it off right away, and I told him I’d be happy to shove my folder of pneumatic rib examples his way, and if he found it useful, I’d be grateful for an acknowledgment. In the actual event, he and Julia asked me to come on as a coauthor, and we were steadily making progress.
That fall I happened to be at Research Casting International at the same time as Brian Curtice — we were both there to see Haplocanthosaurus delfsi while it was down off exhibit from the Cleveland Museum. I’d hung out with Brian a lot back in grad school, but with one thing and another we hadn’t seen each other in many years, and those few days at RCI were a welcome opportunity to rekindle our friendship (and start down the path to coauthorship). Brian also got a look at YPM 1980, the holotype skeleton of Brontosaurus excelsus, while it was at RCI for a remount. Lo and behold, he found unmistakable pneumatic cavities in two of the dorsal ribs of YPM 1980.

A, left rib I, and B, right rib II of YPM 1980, the holotype of Brontosaurus excelsus, in posterior view. King et al. (2024: fig. 2).
That’s pretty awesome for a few reasons. We already knew that the dorsal ribs could be pneumatic in Apatosaurus louisae, because one of the ribs of CM 3018 has a nice round pneumatic cavity. But there was no solid evidence of costal pneumaticity in Brontosaurus. Marsh (1896) figured a rib with pneumatic cavities and claimed it for Brontosaurus, but without a specimen number the referral was uncertain. Turns out there is costal pneumaticity in Brontosaurus, and not just any bronto, but the ur-brontosaur itself, YPM 1980. And in 143 years, no-one had clocked it (there’s a lot of that going around). It seemed silly to write up a pneumatic rib of Apatosaurus from Mygatt-Moore and not mention the newly-discovered rib pneumaticity in YPM 1980, so we brought Brian in on the project. The manuscript went through a genuinely constructive review process at JVP, and we were revising the text and figs last fall.
While I had the apatosaur rib pneumaticity paper with Logan, Julia, and Brian going on one burner, Mike went to Chicago, decided that Brachiosaurus ribs were worth looking at after all (full story here), and went and wrote an entire paper on them in essentially no time. So after deciding in July of 2022 that I was never going to get around to my sauropod rib paper and I should hand it off to someone else (which was absolutely the right decision), a mere 14 months later I found myself working on two sauropod rib papers simultaneously. But they were on different taxa and had somewhat different focuses, so I made my junior author contributions to both and tried not to let Brachiosaurus step on Apatosaurus’s toes. (In particular, Mike and I didn’t talk much about pneumatic ribs outside of Brachiosauridae because there was already a broader survey in Logan’s manuscript.) Brach flew through review and into print just before year’s end (Taylor and Wedel 2023), and now the apatosaurines have lumbered over the finish line. I’m proud of both papers, and very happy to have them out in the world.

Proximal rib head that compromises MWC 9617 in posterior view. The inset image depicts a line drawing of the section of the rib that preserves pneumatic fossae within the rib canal sulcus. Abbreviations: cp, capitulum; I, proximal pneumatic fossa; II, middle pneumatic fossa; III, distal pneumatic fossa; t, tuberculum. Scale bar equals 5 cm. King et al. (2024:fig. 1).
MWC 9617 is an interesting specimen, with a series of same-sized fossae running down the postero-medial side, inside a long sulcus. That’s the side of the rib where the intercostal nerve, artery, and vein would have run — because that’s where they run in all tetrapods — but that neurovascular bundle doesn’t usually sit in a sulcus in sauropod ribs (the same neurovascular bundle does sit in a groove on the underside of human ribs). Those fossae are too smooth and too regular to be pathological. Pneumatic excavations that far down the rib shaft are unusual but not unprecedented — some of the ribs of Paluxysaurus and the Wyoming Supersaurus have pneumaticity about that far distally, and then there’s the weird lonely foramen in the one rib of Brachiosaurus that Riggs (1904) did illustrate. And sometimes pneumatic diverticula do create repeated excavations that look almost identical; one of my favorite examples is the series of pneumatic foramina on the right side of the centrum in a cervical vertebra of (perhaps fittingly) Paluxysaurus. So this certainly looks like a large pneumatic excavation, which we might call a fossa or a sulcus, containing smaller subfossae excavated at regular intervals. That’s pretty cool, because although that general mode of pneumatization turns up now and then in vertebrae, nobody’s documented it in a rib before.

C5? of Paluxysaurus in right lateral view, traced from a photo I took at the Fort Worth Museum of Science and History back in 1990s. I should do a separate post just on this vert sometime — the pneumatic excavations on the left side of the centrum are completely different.
We think that MWC 9617 is a rib of Apatosaurus louisae, for a couple of reasons. One, A. louisae is the most common sauropod at Mygatt-Moore by a wide margin, so any given rib from MMQ is more likely to belong to Apatosaurus than to anything else. The other sauropods known from MMQ so far are Camarasaurus and an indeterminate diplodocine (Foster et al. 2018) — and no pneumatic ribs have ever been described for either Camarasaurus or any of the Morrison diplodocines. (That in itself is pretty weird, given that Diplodocus and especially Barosaurus have pretty complex and extensive vertebral pneumaticity. How did a thicc boi like Apatosaurus beat them to the punch on pneumatizing ribs?) Anyway, it’s more parsimonious that the pneumatic rib from the apatosaur-dominated quarry belongs to Apatosaurus, for which pneumatic ribs are already known, than that it belongs to Camarasaurus or a diplodocine, for which it would be a world first. Bottom line, if we’re wrong, that’s even more exciting.
What’s next? At some point, more stuff from Mygatt-Moore! Jessie and I made Dinosaur Journey home base for our 2022 research trip because neither of us had ever gotten more than one day at a time in that collection. With a whole week to play there, and Julia and Logan to show us weird stuff, we made a LOT of progress, and found some stuff even I didn’t expect. Watch this space.
If you’re around sauropod material, look at ribs. Even the ones that were described in the 1800s may surprise you. Describing pneumaticity is everyone’s business — if you see something, say something!
References
- Foster, J., Hunt-Foster, R., Gorman, M., Trujillo, K., Suarez, C., McHugh, J., Peterson, J., Warnock, J. and Schoenstein, H., 2018. Paleontology, taphonomy, and sedimentology of the Mygatt-Moore Quarry, a large dinosaur bonebed in the Morrison formation, western Colorado—implications for Upper Jurassic dinosaur preservation modes. Geology of the Intermountain West 5:23-93.
- King, J.L., McHugh, J.B., Wedel, M.J., and Curtice, B. 2024. A previously unreported form of dorsal rib pneumaticity in Apatosaurus (Dinosauria: Sauropoda) and implications for pneumatic variation among diplodocid dorsal ribs. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2024.2316665
- Marsh, O.C. 1896. The Dinosaurs of North America. 16th annual report of the U. S. Geological Survey, 1894-95, pt. I. US Government Printing Office, Washington, D.C.
- Riggs, Elmer S. 1904. Structure and relationships of opisthocoelian dinosaurs. Part II, the Brachiosauridae. Field Columbian Museum, Geological Series 2(6):229-247, plus plates LXXI-LXXV.
- Taylor, Michael P., and Mathew J. Wedel. 2023. Novel pneumatic features in the ribs of the sauropod dinosaur Brachiosaurus altithorax. Acta Palaeontologica Polonica 68(4): 709–718. doi:10.4202/app.01105.2023


















