Here at SV-POW!, we love bifurcated cervical ribs. Those of Turiasaurus are one of the autapomorphies proposed by Royo-Torres et al. (2006:figure 1K). Their diagnosis of the new genus included “accessory process projecting caudodorsally from the dorsal margin of the shafts of proximal cervical ribs”. Here is the best example of such a rib in Turiasaurus, attached to its vertebra. (It’s a shame the black backdrop doesn’t extend beneath the rib, but you can make it out easily enough nevertheless.)
This is a very similar but not identical photo to that used in Royo-Torres et al’s (2006:figure 1K) illustration; but because that paper was unfortunately published in Science instead of in a scientific journal, the illustration is microscopic and the description perfunctory. There are no further illustrations of the material in the supplementary document.
Aaanyway. We mentioned but did not illustrate this rib in our recent paper (Wedel and Taylor 2023), and we wrote of it (on page 93) that:
Royo-Torres et al. (2006) described and illustrated bifurcated cervical ribs in Turiasaurus, and Britt et al. (2017) described and illustrated bifurcated cervical ribs in Moabosaurus (Fig. 3A). Turiasaurus and Moabosaurus are both members of the clade Turiasauria, but bifurcated ribs are absent in Mierasaurus, which Royo-Torres et al. (2017) recovered as the sister taxon of Moabosaurus within Turiasauria. This implies either a single origin of bifurcated cervical ribs in Turiasauria, with a reversal in Mierasaurus, or parallel origins of bifurcated cervical ribs in Turiasaurus and Moabosaurus. [emphasis added]
I think we missed the most obvious explanation here: that while the potential to develop bifurcated cervical ribs is phylogenetically determined, the actual development of bifurcation in any given rib is highly variable between individuals and indeed between vertebrae of a single individual. Given that we showed this individual variability pretty clear for apatosaurines — within the Apatosaurus louisae holotype CM 3018, for example — it’s a bit dumb that we failed to apply the same observation to the variable appearance of bifurcated cervical ribs in turiasaurs.
Anyway, that’s not why we’re here. We’re here to look again at how different the bifurcation is between Turiasaurus and some apatosaurines. Here’s a composite, based on the photo above and our old friend MWC 1946 (also appearing as Wedel and Taylor 2023:figure 3d).
(Why don’t we know the specimen number of the Turiasaurus cervical? Again, because the “description” of this important and unusually complete sauropod was published in Science, which is far too important to waste space on trivia like specimen numbers. All we’re told is that the holotype elements have specimen numbers in the range CPT-1195 to CPT-1210. Once, more Science is the opposite of science. Digression ends.)
Back in November last year, Matt asked the question: Single-segment neck muscles in diplodocids?. He tentativly concluded yes, based on the posterodorsal trajectory of the upper prong in apatosaurine cervical ribs, which would have anchored short flexor colli lateralis muscles attaching to the cervical rib loop of the immediate successor vertebra. (We know where on the vertebra these muscles originate in birds, and the upward-and-backward orientation od the prong points to where that site is on the very next vertebra.)
Looking at the composite image above suggests that the same was not true in Turiasaurus. Here, the upper and lower prongs of the bifurcated rib are close to parallel, implying that flexor colli lateralis muscles inserting on the upper prongs can only have originated some segments further back.
Now if we return to the Freak Gallery in our recent paper:
We can see that the rib of Moabasaurus has near-parallel prongs like those of its relative Turiasaurus — implying that it, too, likely had multi-segment flexor colli lateralis muscles. But by contrast the Dicraeosaurus rib, which we described as “incipiently bifurcated”, has a more or less dorsally projecting flange which seems likely to have anchored a single-segment muscle as in apatosaurines.
To my tentative conclusion is that bifurcated ribs in diplodocids (such as apatosaurines and Dicraeosaurus) do indeed imply single-segment neck muscles (or at least single-segment flexor colli lateralis); whereas the bifurcated ribs of turiasaurians imply multi-segment muscles.
Admittedly this is a biggish conclusion to hang on such scant evidence as the apparent angle of divergence in the handful of bifurcated cervical ribs that we’ve seen. But I think it at least stands as the hypothesis best supported by presently available evidence, and it’s there to be corroborated or contradicted by further observations.
References
- Royo-Torres, Rafael, Alberto Cobos and Luis Alcalá. 2006. A giant European dinosaur and a new sauropod clade. Science 314:1925–1927.
- Wedel, Mathew J., and Michael P. Taylor. 2023. The biomechanical significance of bifurcated cervical ribs in apatosaurine sauropods. VAMP (Vertebrate Anatomy Morphology Palaeontology) 11:91-100. doi: 10.18435/vamp29394
As iconic as Brachiosaurus altithorax is, it’s known from surprisingly little material. As I cover in my 2009 brachiosaur paper (Taylor 2009:788–789), only the holotype specimen FMNH PR 25017 can be reliably considered to belong to the species: little of the material that has been referred to it over the years overlaps with the holotype, and among those elements that do, synapomorpies are hard to come by.
One referred element that does overlap is the Potter Creek humerus, which I covered in that paper as follows (Taylor 2009:788):
Potter Creek Humerus—As recounted by Jensen (1985, 1987), Eddie and Vivian Jones collected a large left humerus from the Uncompahgre Upwarp of Colorado and donated it to the Smithsonian Institution where it is accessioned as USNM 21903. It was designated Brachiosaurus (Anonymous, 1959) although no reason for this assignment was published; it was subsequently described very briefly and inadequately by Jensen (1987:606-607). Although its great length of 213 cm (pers. obs.) is compatible with a brachiosaurid identity, it is in some other respects different from the humeri of both B. altithorax and B. brancai, although some of these differences may be due to errors in the significant restoration that this element has undergone. The bone may well represent Brachiosaurus altithorax, but cannot be confidently referred to this species, in part because its true proportions are concealed by restoration (Wedel and Taylor, in prep.). It can therefore be discounted in terms of contributing to an understanding of the relationship between B. altithorax and B. brancai.
(By the way: that Wedel and Taylor (in prep.) paper has not materialized, fifteen years on. It’s titled “The humeri of brachiosaurid sauropods” and the manuscript has not been touched since 2007 — two years before the main brachiosaur paper was published! I just looked at it, and it’s 14 pages long, so I guess that’s yet another project that we really ought to exhume and push over the line.)
I first encountered this humerus in Jensen, where it’s illustrated in Figure 4:
Obviously you can’t make out a ton of detail in this photo, which in any case is of a replica rather than the original bone. But Jensen illustrated it better in his 1987 paper, figures 3 and 5 (as well as repeating figure 4 of his 1985 paper as figure 6 of the 1987 one).
Jensen’s caption doesn’t say it, but obviously this view is anterior. (The dorsal vertebra from the same quarry is a whole nother kettle of non-tetrapod vertebrates, which we won’t discuss today.)
This is not the clearest illustration. Part A is obviously in anterior view, matching nicely with Jensen’s figure 3E. Part B seems to be in medial view, and part C in lateral view. Part D, I can’t make much sense of: it’s described as “anterior, distal end”, but it’s not a good match for the distal end shown in figure 3E.
Some time later, I got to see the bone for myself: it’s long been on public display as a touch specimen at the NMNH in Washington DC. Here’s a photo — not one of mine, which didn’t come out too well, but one sent by Mike Brett-Surman:
Now you can probably tell from the photo, but in person it was really obvious that a great chunk in the middle was fakezilla. Here’s the drawing I did for myself back in 2007:
(Yes, my sketch has the proportions horribly wrong. But it does properly capture where the faked up areas are.)
And here is one of my not-very-good photos: a close-up of part of the shaft, where damage to the surface clearly shows that what’s underneath the gloss is not bone but fibreglass or something similar:
I think that reconstructed shaft is wider than it should be, which is why I argued back in 2009 that “it is […] different from the humeri of both B. altithorax and B. brancai, although some of these differences may be due to errors in the significant restoration that this element has undergone […] its true proportions are concealed by restoration.”
I’d since come around to thinking the humerus most likely is Brachiosaurus after all, as the main reason for finding that unlikely is down to the reconstructed thick shaft. But a little while ago, I found something really helpful: Brachiosaurus photos in the Smithsonian Institution Archives! In particular, this one showing the humerus as it used to be in 1959, shortly after it was donated to the museum sitting on a plinth in front of the mounted Diplodocus forefeet (which are really Camarasaurus forefeet, but that’s a different story).
The exciting thing about this photo is of course that it was taken before all that midshaft restoration was done. And sure enough, the shaft is noticeably narrower than in the current restored version — a much better match for the holotype Brachiosaurus humerus and even the yet-more-slender one of Giraffatitan.
(There is more I could say here, notably about the deltopectoral crest. But that can wait for another day — this post is plenty long enough as it.)
I can understand why this restoration was done: if this was to be a touch specimen, that fragile, damaged mishaft was absolutely going to flake away. The purpose of the restoration was probably just protection. But I still lament that it was done — and that it was done in this way. To me, it just says that this should never have been a touch specimen.
References
- Anonymous. 1959. Brachiosaurus exhibit at the Smithsonian Institution. Nature 183:649–650.
- Jensen, James A. 1985. Three new sauropod dinosaurs from the Upper Jurassic of Colorado. Great Basin Naturalist 45(4):697-709.
- Jensen, James A. 1987. New brachiosaur material from the Late Jurassic of Utah and Colorado. Great Basin Naturalist 47(4):592-608.
- Taylor, Michael P. 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914). Journal of Vertebrate Paleontology 29(3):787-806. doi: 10.1671/039.029.0309
I was cleaning out my Downloads directory — which, even after my initial forays, still accounts for 11 Gb that I really need to reclaim from my perptually almost-full SSD. And I found this beautiful image under the filename csgeo4028.jpeg.
The thing is, I have no idea where this image came from. The file’s timestamp says it’s been 16 months since I downloaded it from somewhere, but there is no associated metadata that tells me where I got it. Googling for the filename gets me nothing.
Can anyone find the source? [Update: see the first coment below! Crown House found it in the Field Museum’s own Geology Collections.]
Anyway, I immediately recognised it as our old friend Brachiosaurus altithorax, and in fact it’s a much better version of a photo that we’ve featured here before, That version was scanned from Supplement 1 of Don Glut’s encyclopedia, which credits it as being Field Museum neg. #4027 (which is one out from the number in the filename). But that doesn’t explain where this high-resolution copy came from.
Anyway, looking at this image in 2024, I’m immediately interested in the ribs, which of course Matt and I published on at the very end of 2023 (Taylor and Wedel 2023, natch). It shows both ribs A and B in their original state, and it’s instructive to compare them with those ribs as we illustrated them in our paper.
First, rib A:
This one is visible at the bottom of the photo, proximal end to the bottom, but flipped over from the way it now rests in the collection, so the pneumatic opening is not apparent. There’s an interesting “folded over” ridge running down the anterior(*) face of the proximal part of the shaft.
(*) Assuming we were right in interpreting the available face of the rib as posterior in our paper.
Now, Rib B:
This is visible on the left side of the image, close to the vertebral column, with the proximal end to the top. It has the same (posterior, we think) face upwards as is available in the collection, and you can make out the pneumatic opening in the tuberculum that we illustrated.
Reference
Eighteen months ago, I noted that the Carnegie Museum’s Diplodocus mount has no atlantal ribs (i.e. ribs of the first cervical vertebra, the atlas). But that the Paris cast has long atlantal ribs — so long the extend past the posterior end of the axis.
There were two especially provocative comments to that post. First, Konstantin linked to a photo of the Russian cast (first mounted in St. Petersburg but currently residing in Moscow). I’ll reproduce it here:
As you can see, there are atlantal ribs on this specimen, but they do not resemble those of the Paris cast. These are much shorter, narrower, and lacking in structure. I have not to my knowledge seen anything like this on any other Diplodocus, and my guess — it’s only a guess — is that these were added by the Russians at some stage in this specimen’s very complex history.
But wait, there’s more!
In another comment on the same post, Crown House linked to a 3D model of the Vienna cast that has been posted to Sketchfab. It’s a pretty low resolution model, but if you zoom and pan, you can see that it has large and complex Paris-style atlantal ribs:
Although these resemble the atlantal ribs of the Paris mount, they are not identical: the wavy margins face posteriorly rather than anteriorly as in both the Paris mount and Holland’s (1906) illustrations; and the proximal end has a dorsal expansion.
So we seem to have (at least) four different state of atlantal ribs in different casts of the same Diplodocus:
- Absent in Pittsburgh and London
- Small and rod-like in Moscow
- Long with a wavy dorsal margin in Paris
- Long with a wavy ventral margin and a proximal dorsal expansion in Vienna
Can anyone offer informed speculation on how this state of affairs came about?
But then things get weird. If you manoeuvre your way around the model to look up at this region from below:
Well, what the heck are we seeing here? There are two spiny processes, one on each side, projecting laterally from the ventral part of the atlas, and swept back at mid-length.
I have never seen anything like this in any sauropod — or, come to think of it, any other animal, but I admit I don’t pay much attention to other animals.
Does anyone have any idea what these projections are? Remember you can go to the model and look at them in 3D.
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.
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.
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.
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
This is one of those things that has been sitting in my brain, gradually heating up and getting denser, until it achieved criticality, melted down my spinal cord, and rocketed out my fingers and through the keyboard. Stand by for caffeine-fueled testifyin’ mode.
Part 1: Why Study Pneumaticity
Last item first: why you should study pneumaticity. The honest reason that primarily motivates me is that pneumaticity is frickin’ cool. Air inside bones! And endlessly novel — pneumatization is opportunistic and invasive (Witmer 1997), and it never quite works out the same way twice. So every time I see a pneumatic bone, inside or out, my antennae are up, because I suspect it will have its own little quirks and oddities, any one of which might unlock something new about the morphogenetic process of pneumatization or its functional importance.
If you need something more respectable than “Whoa, dude!” to put on a thesis proposal or a grant application, how’s this: we think that skeletal pneumaticity was a key innovation for both sauropods (Sander et al. 2011) and theropods (Benson et al. 2012) [edit: and pterosaurs {Claessens et al. 2009}], but our documentation of it is very poor. For a lot of sauropod genera, we’ve only CT-scanned one or two vertebrae, often from the same quarry, usually from a single individual. For a lot more, we’ve scanned none at all. As I wrote back in 2018, “Someone just needs to sit down with a reasonably complete, well-preserved series that includes posterior dorsals, all the sacrals, and the proximal caudals–or ideally several such series–and trace out all of the pneumatic features” (link). The same principle — “crawling” one or more specimens to document everything — could be extended to address intraspecific and interspecific variation, the extent to which pneumatic traces might relate to nerve and blood vessel pathways, and ontogenetic changes. We know that vertebral pneumatization got more extensive and more complex through an individual animal’s maturation, but we don’t know much about how and when that happened, or if it ever stopped in large and long-lived individuals. I don’t know what we’ll find when people get around to doing this, but there won’t be any boring answers — indeed, much of what I thought about the early evolution of pneumaticity for the last 25 years is probably wrong.
Whether you want to work on pneumaticity or not, definitely do not make the mistake of looking at the existing literature and assuming “it’s all been done“. I’ve probably spilled more ink about dinosaur pneumaticity than anyone else alive, and I’m telling you that the field is wide open. Just off the top of my head:
- Sometimes pneumatized sauropod vertebrae have more bone than they need, because fossae are embossed into otherwise flat plates of bone that would be lighter if they lacked those fossae. What’s up with that? Does it ever happen in theropods (avian or otherwise) or pterosaurs?
- I mentioned that pneumatic bones rarely look identical under the hood. Heck, they rarely look identical on the surface. Whether it’s internal or external asymmetry, or variable laminae, or some other thing, there’s a LOT of variation. How does that small-scale morphogenetic opportunism jibe with the apparent macroevolutionary importance of pneumaticity in sauropods and theropods [edit: and pterosaurs]?
- Related: my a priori assumption is that pneumaticity was functionally important in non-avian theropods, more functionally important in sauropods (because size), and most functionally important in pterosaurs (because size x flight). That’s a wild guess, totally untested — but I’ll bet someone will figure out a way to test it, and variation vs developmental constraint seems like fertile ground for that testing.
- Also related: does skeletal asymmetry (pneumatic or otherwise) have any predictable relationship with body size, either ontogenetically or phylogenetically? See this post and this one for some related noodling (but no answers).
- For internal pneumatization, do bigger and older individuals make more chambers that are about the same size as the chambers in smaller individuals, or does the subadult level of complexity stay the same through adulthood, and the chambers get bigger but not more numerous? And is there even a single answer, or do different things happen in different lineages? These seem like fundamental questions, and I have my suspicions, but AFAIK neither I nor anyone else has addressed this. Put a pin this, it will come up again later in this post.
- Barosaurus cervicals have a more complex internal structure than Diplodocus or Apatosaurus cervicals (check out the eroded condyle of this vertebra). Is that because Barosaurus cervicals are longer? Is there a functional reason we never see crazy long vertebral centra that are camerate?
- Want to work on birds? Do some injections and dissections and see how often diverticula follow nerves and blood vessels as they develop. This idea, which has a lot of circumstantial support (Taylor and Wedel 2021), is based on a single observation from a paper published nearly a century ago (Bremer 1940).
- Heck, if you’re doing injections and dissections, just document the diverticular network in a single bird, full stop. That’s a descriptive paper right there. Bird pneumaticity is so grossly understudied that whole classes of diverticula are still being described for the first time (Atterholt and Wedel 2022).
- Rather work on sauropods or non-avian theropods? We could use a lot more work on pneumosteum (Lambertz et al. 2018), and on the histological signals of pneumaticity, in basically everything from pig sinuses to the tail of Diplodocus — especially basal sauropodomorphs and early theropods where pneumaticity was just getting up and running.
- Don’t want to do histo? CT scan something. Anything. And write it up. Especially dorsals, sacrals, and caudals — the published sample is skewed toward cervicals because they’re long and skinny and fit through the machines better. Don’t have access to a CT machine? No worries, that’s what the second half of this post is about.
- Don’t want to mess with machines at all? Crawl some skeletons — or maybe just like one fairly complete diplodocid or titanosaur — and describe the pneumatic (and maybe also vascular) features on the ventral surfaces of the vertebrae. That’s a whole class of diverticula (or maybe multiple classes) about which we know basically zip, other than that sometimes cervicals and caudals have foramina on their ventral surfaces (but not dorsals or sacrals — why?). You might be able to get a short review paper just canvasing examples in the literature — but if you don’t go look at specimens in person, you’ll miss a lot, because these features are are rarely described or illustrated.
- Want a project you can do on the couch in your jammies? Wedel (2003) is my most-cited paper by some distance, but it’s waaay out of date. Comb the literature and write an up-to-date version of that paper just based on all the new stuff that’s been published in the past two decades. Here’s a fun starter: I made a big deal in that paper about camerate vertebrae in a then-undescribed titanosaur from Dalton Wells in the Cedar Mountain Formation. In time that critter proved to be Moabosaurus, a turiasaur and not a titanosaur. The whole idea of camerate titanosaurs needs a re-look. And I didn’t write anything about turiasaurs back then because the clade hadn’t been recognized yet. My top paper, and at this point it might as well have been scratched out on clay tablets. (Note: this is a good thing. That paper is out of date because there’s been so much progress. If it was still cutting-edge, it would mean the field of sauropod pneumaticity was dead. But still — someone go knock that thing off its perch.)
How to Study Pneumaticity on the Cheap
I think there is an assumption, or a perception, that you need to CT scan fossils to study pneumaticity. Access to CT scanners can be logistically complex, and expensive. Can be, not has to be. And there’s a lot of crucial work to be done without a CT machine. Let’s get to it.
1. Collaborate with a radiologist. Okay, but what if you do want to CT scan some fossils? Do what I do, and ask around to see if there’s a radiologist who is interested in collaborating. Most hospital CT machines are not busy all the time — there’s usually one slow afternoon each week, or each month. And in my experience, most radiologists are down to look at something interesting and different, like a dinosaur bone, as a break from the endless parade of concussions, degenerated lumbar discs, and cirrhotic livers. The collaboration piece is key. I’m not a radiologist, and minimally I need a professional who can write up the machine specs and scan settings for the Materials and Methods section of the paper. But often the radiologist will see interesting things in the scan that I would have missed, or I’ll see interesting things in the scans that may turn out to be mundane features that look weird in cross-section. And I’m more than happy to trade authorship on whatever papers come out of the scans, and acknowledgement and good press for the hospital, in exchange for the professional’s expertise and time on the machines. Specific advice? Be humble, be polite. Once I’m through the hospital doors I’m not the expert in anything other than safely handling the fossils, and I make it clear that I’m there to be safe, respect their turf, let them direct the logistics, and learn as much as I can. All the radiologists I’ve worked with have been happy to share their knowledge, and curious about the fossils and what we hope to learn from the scans.
2. Use broken specimens. I’ve blogged before about how breaks and erosion are nature’s CT machines (here, here, here, and here, for starters), and I’ve favorably discussed the utility of broken specimens in my papers, but I figured broken specimens would always be distant also-rans in the quest to document pneumaticity. Then I read Fronimos (2023) — hoo boy. John Fronimos set out to document pneumaticity in a Late Cretaceous titanosaur from Texas (maybe Alamosaurus, maybe not), and he crushed it. It’s one of the best danged sauropod pneumaticity papers I’ve ever read, period, and the fact that he did it all without CT scanning anything makes it all the more impressive. And it’s not only a great descriptive paper — John’s thoughts on the evolution and function of pneumaticity in sauropods are comprehensive, detailed, insightful, and forward-looking. Up above I mentioned reading broadly to get caught up; if you work on sauropod pneumaticity, or want to, or just want to understand the state of the art, the discussion section of Fronimos (2023) is the new bleeding edge. Also, remember the pin we placed up above, on the question of whether pneumatic chambers get bigger or more numerous or both over ontogeny? With the right collection you could answer that with only broken specimens.
3. Study external pneumatic features. This has already come up a few times in this post, but let me draw the threads together here. Whether it’s documenting serial changes in pneumatization along the vertebral column in a single individual, or externally-visible asymmetry, or pneumaticity on the ventral surfaces of vertebrae, or how and whether pneumatic and neurovascular features relate to each other, there is a ton of work to be done that just requires collections access, a notebook, a camera, and time. And it lends itself to collaboration; two sets of eyes will see a lot more. (If you have the freedom to choose, ideally you might want one fairly big and strong person to manhandle the bones [safely, for the sake of the bones and the humans], and one fairly slim and flexible person to scramble up ladders and fit into odd nooks and crannies.)
4. Use publicly-available CT data. Okay, admittedly there’s probably not enough of this out there yet to use on anything other than birds (or mammals, if you’re into sinuses), but hey, we need bird studies, too. Bird studies hit twice — first because birds are interesting objects of study in their own right, and second because they’re our baseline for interpreting pneumaticity in fossils. (By quick count, I’ve figured drawings, photos, or CT scans of bird vertebrae in more than dozen of my papers, and in half a dozen cases they were vertebrae I prepped myself at home.) Of the four paths, this is the one I have the least experience with, but the new “oVert” (openVertebrate) collection on MorphoSource is a good place to start. Wet specimens may have a bit of a learning curve in terms of distinguishing pneumatic and non-pneumatic bones, and most of the extra-osseous pneumatic diverticula have probably collapsed, but with free access to CT scans of “>13,000 fluid-preserved specimens representing >80% of the living genera of vertebrates” I’ll bet people will think of plenty of cool stuff to do. Here’s the oVert trailer:
Conclusion: Let’s Roll
We need more pneumaticity studies. There is just so much we don’t know. I’ve been working on sauropod pneumaticity more often than not since 1998, and I’m stoked about how much basic descriptive work remains to be done, because I’m an anatomy geek at heart, and describing weird anatomy is deeply satisfying for me, as is reading other people’s descriptions of weird anatomy. But I’m also in despair about how much basic descriptive work remains to be done, because the answers to so many questions are still over the horizon from us, and probably will be for the rest of my life.
So please, if you’re interested, come do this work. Whether you’re a grad student at a major institution with an NSF pre-doc fellowship and several years of runway in which to do unfettered research, or just some person sitting on a couch thinking about dinosaur bones (er, like me right now), now you have some ideas to work on (or reach beyond), and some inexpensive ways to work on them. If you’re curious and want to get your feet wet before you commit, remember that you can get extant dinosaur carcasses at the grocery store, and prep and section your own pneumatic dinosaur bones at the kitchen table. There is a very accessible on-ramp here for anyone who has the time and inclination. Let’s do this thing.
References
- Atterholt, Jessie, and Wedel, Mathew J. 2022. A computed tomography-based survey of paramedullary diverticula in extant Aves. The Anatomical Record, 1– 22. https://doi.org/10.1002/ar.24923
- Benson, R.B., Butler, R.J., Carrano, M.T. and O’Connor, P.M., 2012. Air‐filled postcranial bones in theropod dinosaurs: physiological implications and the ‘reptile’–bird transition. Biological Reviews, 87(1), pp.168-193.
- Bremer, John L. 1940 The pneumatization of the humerus in the common fowl and the associated activity of theelin. The Anatomical Record 77(2):197–211. doi:10.1002/ar.1090770209
- Claessens LPAM, O’Connor PM, Unwin DM (2009) Respiratory evolution facilitated the origin of pterosaur flight and aerial gigantism. PLoS ONE 4(2): e4497. doi:10.1371/journal.pone.0004497
- 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
- Lambertz, M., Bertozzo, F. and Sander, P.M. 2018. Bone histological correlates for air sacs and their implications for understanding the origin of the dinosaurian respiratory system. Biology Letters 14(1): 20170514.
- 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.
- Taylor, Michael P., and Mathew J. Wedel. 2021. Why is vertebral pneumaticity in sauropod dinosaurs so variable? Qeios 1G6J3Q. doi:10.32388/1G6J3Q
- Wedel, M.J. 2003. The evolution of vertebral pneumaticity in sauropod dinosaurs. Journal of Vertebrate Paleontology 23:344-357.
- Witmer, L.M. 1997. The evolution of the antorbital cavity of archosaurs: a study in soft-tissue reconstruction in the fossil record with an analysis of the function of pneumaticity. Journal of Vertebrate Paleontology 17(Supplement 1): 1-76.
Tutorial 43: how to do creative work
March 19, 2024
I was struck by a Mastodon post where classic game developer Ron Gilbert quoted film critic Roger Ebert as follows:
The Muse visits during the act of creation, not before. Don’t wait for her.
And Gilbert commented:
I am constantly forgetting this as I procrastinate writing only to discover her again once I start.
In a reply, Gretchen Anderson said her favourite version of this is:
When the muse comes to visit, she better find you working.
I couldn’t find the original source for this, but as I was trying to track it down I ran into this, attributed to Pablo Picasso:
Inspiration exists, but it has to find you working.
When I mentioned these observations to Matt, he sent me a longer-form exposition of the same phenomenon, from painter and visual artist Chuck Close. If the four pithy versions above strike you as a bit facile, the kind of thing you might find on a motivational poster, this may be more actionable:
All the best ideas come out of the process; they come out of the work itself. Things occur to you. If you’re sitting around trying to dream up a great idea, you can sit there a long time before anything happens. But if you just get to work, something will occur to you and something else will occur to you and something else that you reject will push you in another direction. Inspiration is absolutely unnecessary and somehow deceptive. You feel like you need this great idea before you can get down to work, and I find that’s almost never the case.
All of this points back, of course, to the very snappiest version of this observation, attributed to William Wordworth and appearing in the previous tutorial:
To begin, begin.
Why should this be true? I don’t know why, but I know that it is. My best guess is that it’s the same principle by which you can only steer a moving boat. Without some kind of resistance to push against, your rudder can’t exert a useful force; without some raw material for it to work on, your creativity can’t get a toehold. You can’t be creative about nothing. You need something to be creative about.
So the substance of this tutorial is this: whatever project you’re procrastinating on — the novel that you can’t quite figure out the ending of, the paper on a subject that you’ve not read enough background information on, the song that you can’t bring yourself to write because it won’t be as good as a Paul Simon song[*] — just make a start. The worst-case scenario is that you’ll better understand where the real blockages are. A better case is that you’ll find there aren’t any. And the best case is that the very process of getting your hands dirty leads you to fresh new ideas.
I leave you with this brief reminiscence from Greg Gunther, who ran a mailing-list I used to be on for aspiring writers:
I was on an [email] list with Tom Clancy once. Mr. Clancy’s contribution to the list was, “Write the damn book”.
(I am pleased to see that we’ve quoted this at least twice before in SV-POW! tutorials: in my own Tutorial 10: how to become a palaeontologist; and in Matt’s Tutorial 12: How to find problems to work on. It remains by some distance the very best advice I’ve ever heard on writing.)
[*] This last one is very much aimed at myself.
On the tragic fate of PeerJ
March 17, 2024
I said last time that Jisc’s feeble transition-to-open-access report was the first of two disapointing scholarly-communication announcements that week. The second was of course the announcement that PeerJ has been acquired by Taylor and Francis.
Matt and I have both been big fans of PeerJ since before it launched, and we were delighted to have our 2013 neck-anatomy paper in the first batch of articles published there. We’ve had a lot of good things to say about its open peer-review, its usefulness in teaching, its disruptiveness, about how it became our default choice of venue, and much more.
So it’s tragic to see it being eaten by one of the legacy publishers.
What’s even more tragic is to see the founders, who I have liked and respected for more than a decade, spouting such transparent b.s. in the press release:
“Becoming part of Taylor & Francis is an important step in PeerJ’s evolution,” explained Peter Binfield, PeerJ Co-Founder and Publisher. “This move will allow us to cement our original commitments to open research, equitable and inclusive publishing and rigorous peer review.”
Jason Hoyt, PeerJ Co-Founder and CEO added: “Our mission to make scientific research accessible to all whilst delivering 21st century technology aligns perfectly with Taylor & Francis’ vision.”
None of this is true. We know it’s not true. Pete and Jason know it’s not true. We know they know it’s not true. They know that we know they know. Why even insult us with this nonsense?
I suppose it’s part of the contract they signed with their new bosses, that they have to make public statements about how excited they are. But, seriously, who is buying this?
Here are two good things about the situation, though:
First, because everything published by PeerJ has been under the Creative Commons Attribution (CC By) licence, all the papers are still and will always remain free to read, redistribute, modify, etc. That’s the wonder of a licence. It’s not dependent on anyone’s good will. There’s nothing Taylor & Francis can do to change this. Even if they were to change PeerJ to use a different and more restrictive licence for new papers (and there’s no reason to think they would), nothing can change the legal status of what’s already been published.
And second, an email sent to PeerJ members promises that the publish-forever deals that we bought back in 2013 are still good:
You may have already seen the recent announcement that PeerJ has been acquired by Taylor & Francis, and I wanted to provide you with some reassurance regarding the status of your PeerJ Lifetime Membership: despite this change, your Lifetime Membership with PeerJ remains valid.
Your commitment to PeerJ and support for our mission are greatly appreciated and valued by PeerJ. If you have any questions or concerns about your lifetime membership or the acquisition in general, please don’t hesitate to reach out to us directly (communities@peerj.com). Your peace of mind is important to us, and we are here to address any inquiries you may have. You can read more in the announcement here.
I wonder how long Taylor and Francis will continue to honour this promise, though. Forever, as initially promised? Ten years? Five? We’ll see what happens when integrity runs up against profit margins.
I’ll have more to say about this acquisition, and about what it tells us about the scholarly publishing landscape more generally, but I’ll leave it there for now. Bottom line for me: this is a very sad day.
Jisc should not buy services that it does not want
March 15, 2024
In the first of two disapointing scholarly-communication announcements last week, Jisc announced its report on progress towards open access in the UK. The key finding is:
Despite improvements – rapid growth in transitional agreements, sector savings and high levels of funder compliance – a full transition to open access will not happen soon.
But that’s not the part that disappoints me. Here’s the part that disappoints me:
Transitional agreements were envisioned as a temporary mechanism to support a transition to full open access. They set out to constrain costs for institutions and drive rapid growth of open access at a UK and global level.
However, the review suggests they are at risk of becoming the ‘norm’ and that the rate of transition is too slow.
Sometimes I think people don’t know what “transitional” means. Folks, a transitional agreement is by definition one that gets you from one place to another — in this case, from a subscription-based scholarly publishing ecosystem to an open-access one. It’s what a transitional agreement is for. If it does not do that, then it’s not transitional. In particular, a transitional agreement that becomes “the ‘norm'” is a stalled agreement. It is, unambiguously, a failure.
So that was the part that disappointed me. And here comes the part that infuriates me (quoted from Professor Stephen Decent, chair of the Jisc UUK Content Negotiation Strategy Group):
Despite significant sector-wide investment, the transition from the paywall system to full open access remains elusive. It’s clear that our open research scholarship objectives are out of step with many publishers’ commercial strategies.
Why does Jisc give a nematode’s anal sphincter about publishers’ commercial strategies? Has it forgotten that it’s the customer here? Jisc is not beholden to the preferences of publishers, publishers are required (or, in a sane world, would be) to provide what Jisc wants.
Look. If I go into Tesco to buy a cabbage, it may happen that I find that my cabbage-purchasing objectives are out of step with Tesco’s commercial strategy. But if that happens, I go and buy my cabbage from Sainsbury’s instead.
What I don’t do is buy a Tesco bath-towel instead, because selling bath-towels is their commercial strategy. Why not? Because I want a cabbage, not a bath towel.
I trust I do not need to spell this out, but I will anyway for avoidance of doubt: Jisc should not buy what these publishers are selling, but should instead find publishers that are selling what Jisc want to buy.
Why were prehistoric sea creatures so tiny?
March 12, 2024
My friend Toby Lowther wrote to me back in December to ask this question:
As far as I understand it, the general rule for extant species is that it’s much easier to get much bigger underwater than on land, due to the role that water plays in supporting large bodies. But as was recently pointed out to me by a friend, sauropods seem to be much, much larger than any known or recorded prehistoric sea creatures — and, in fact, despite sauropods being much larger than any modern megafauna, plesiosaurs, mosasaurs, etc. seem to be smaller than, say, blue whales. Is there a current theory/explanation as to why? Why you have these various periods where land megafauna ends up larger than any contemporaneous sea megafauna, and why despite dinosaurs getting so big, the largest prehistoric sea megafauna are smaller than the largest modern whales?
It’s strange, isn’t it? The last I knew, Shonisaurus was the largest ichthyosaur, at about 20 m and 50 tonnes, and this is considerably bigger than any plesiosaur or mosasaur I know of. It’s up the sperm-whale size category, but not even close to the bigger baleen whales. Why not?
Meanwhile, there is solid evidence for sauropods massing more than this on land — and less solid but I think still good evidence for them getting at least twice this heavy.
So what’s the deal with all those Mesozoic sea creatures being so tichy compared with terrestrial beasts that had so many more size-related problems to deal with?
Speak, O commenters! We’re listening.