Here’s the story of my fascination with supramedullary airways over the last 20 years, and how Jessie Atterholt and I ended up working on them together, culminating with her talk at SVPCA last week. (Just here for the preprint link? Here you go.)

Müller (1908: fig. 12). Upper respiratory tract, trachea, and lungs in pink, air sacs and diverticula in blue. DSPM = diverticulum supramedullare.

Way back when I was working on my Master’s thesis at the University of Oklahoma and getting into pneumaticity for the first time, Kent Sanders found Müller (1908) and gave me a photocopy. This would have been the spring or summer of 1998, because we used some of Müller’s illustrations in our poster for SVP that year (Wedel and Sanders 1998). Müller’s description of pneumatic diverticula in the pigeon formed part of my intellectual bedrock, and I’ve referenced it a lot in my pneumaticity papers (complete list here).

One of the systems that Müller described is the diverticulum supramedullare, a.k.a. supramedullary diverticula, or, informally, supramedullary airways (SMAs). Traditionally these are defined as pneumatic diverticula that enter the neural canal and lie dorsal (supra) to the spinal cord (medulla), although O’Connor (2006) noted that in some cases the diverticula could completely envelop the spinal cord in a tube of air. I yapped about SMAs a bit in this post, and they’re flagged in almost every ostrich CT or dissection photo I’ve ever published, here on the blog or in a paper.

CT sections of a Giraffatitan cervical, with connections between the neural canal and pneumatic chambers in the spine highlighted in blue. Modified from Schwarz & Fritsch (2004: fig. 4).

Fast forward to 2006, when Daniela Schwarz and Guido Fritsch documented pneumatic foramina in the roof of the neural canal in cervical vertebrae of Giraffatitan. As far as I know, this was the first published demonstration of SMAs in a non-bird, or in any extinct animal. Lemme repeat that: Daniela Schwarz found these first!

OMNH 60718: too ugly for radio. This is an unfused neural arch in ventral view. Anterior is to the left. Neurocentral joint surfaces are drawn over with ladders; pneumatic foramina lie between them.

Shortly thereafter I independently found evidence of SMAs in a sauropod, in the form of multiple pneumatic foramina in the roof of the neural canal in an unfused neural arch of a basal titanosauriform (probably a brachiosaurid) from the Cloverly Formation of Montana. It’s a pretty roadkilled specimen and I was busy with other things so I didn’t get around to writing it up, but I didn’t forget about it, either (I rarely forget about stuff like this).

Then in 2013 I went to the Perot Museum in Dallas to see the giant Alamosaurus cervical series, and I also visited the off-site research facility where juvenile Alamosaurus from Big Bend is housed. When Ron Tykoski let me into the collections room, I was literally walking through the door for the first time when I exclaimed, “Holy crap!” I had spotted an unfused neural arch of a juvenile Alamosaurus on a shelf across the room, with complex pneumatic sculpting all over the roof of the neural canal.

Title slide for the 2014 SVPCA presentation.

The Big Bend and Cloverly specimens were the basis for my talk on SMAs at SVPCA in 2014, coauthored with Anthony Fiorillo, Des Maxwell, and Ron Tykoski. As prep for that talk, I visited the ornithology collections at the Natural History Museum of Los Angeles County, photographed a lot of bird vertebrae with foramina inside their neural canals, and shot this pelican video. That was four years ago – why no paper yet? It’s because I wanted one more piece of smoking-gun evidence: a CT scan of a bird that would show a direct communication between the SMAs and the air spaces inside a vertebra, through one or more foramina in the roof, wall, or floor of the neural canal.

A spectrum of pneumatic traces in the neural canals of birds, including complexes of large or small foramina, isolated foramina, and sculpting without foramina.

In 2017, Jessie Atterholt taught in our summer anatomy course at WesternU as an adjunct (her full-time employment was at the Webb Schools in Claremont, home of the Alf Museum). Jessie and I had been acquainted for a few years, but we’d never had the opportunity to really talk science. As we chatted between dissections, I learned that she had a huge warchest of CT scans of whole birds from her dissertation work at Berkeley (we’d missed each other by a few years). My antennae twitched: one nice thing about SMAs is that, being bounded by bone, they can’t collapse after death, unlike more peripheral diverticula. And air is jet black on CT scans, so SMAs are easy to spot even on comparatively low-res scans. All you need is one or two black pixels. I proposed a collaboration: we could use her CT scans to survey the presence and distribution of SMAs in as many birds as possible.

Vertebral diverticula in two sagittally-exploded cervical vertebrae of a turkey. Anterior is to the left, #5 is the SMA. Cover (1953: fig. 2). Yes, I know this is gross – if anyone has a cleaner scan, I’m interested.

You might think that such a survey would have been done ages ago, but it’s not the case. A few authors have mentioned supramedullary airways, and O’Connor (2006) gave a good description of some of the variation in SMAs in extant birds as a whole. But the only detailed accounts to illustrate the morphology and extent of the SMAs in a single species are Müller (1908) on the common pigeon and Cover (1953) on the domestic turkey. I’d seen what I suspected were traces of SMAs in the vertebrae of many, mostly large-bodied birds, and I’d seen them in CTs of ostriches and hummingbirds, and in ostriches and turkeys in dissection. But Jessie was offering the chance to see both the SMAs and their osteological traces in dozens of species from across the avian tree.

SMAs in a micro-CT of a female Anna’s hummingbird, Calypte anna. Scale bars are in mm.

Real life intervened: we were both so busy teaching last fall that we didn’t get rolling until just before the holidays. But the project gradually built up steam over the course of 2018. One story that will require more unpacking later: everything I’ve written on this blog about neural canals, Haplocanthosaurus, or CT scanning in 2018 is something serendipitously spun out of the SMA survey with Jessie. Expect a lot more Atterholt and Wedel joints in the near future – and one Atterholt et al. (minus Wedel) even sooner, that is going to be big news. Watch this space.

It didn’t hurt that in the meantime Jessie got a tenure-track job teaching human anatomy at WesternU, to run the same course she’d taught in as an adjunct last year, and started here at the beginning of June. By that time we had an abstract on our findings ready to go for this year’s SVP meeting. Alas, it was not to be: we were out in the field this summer when we learned that our abstract had been rejected. (I have no idea why; we’ve increased the taxonomic sampling of SMAs in extant birds by a factor of six or so, most of our important findings are in the abstract, and we mentioned the relevance to fossils. But whatever.)

We were bummed for a day, and then Jessie decided that she’d submit the abstract to SVPCA, only slightly chopped for length, and go to Manchester to present if it was accepted – which it was. Unfortunately I’d already made other plans for the fall, so I missed the fun. Fortunately the SVPCA talks were livestreamed, so last Friday at 1:30 in the morning I got to watch Jessie give the talk. I wish the talks had been recorded, because she knocked it out of the park.

Title slide for the 2018 SVPCA presentation.

And now everything we’re in a position to share is freely available at PeerJ. The SVPCA abstract is up as a PeerJ preprint (Atterholt and Wedel 2018), the longer, rejected SVP abstract is up as a supplementary file (because it has a crucial paragraph of results we had to cut to make the length requirement for SVPCA, and because why not), and our slideshow is up now, too. I say ‘our’ slideshow but it’s really Jessie’s – she built it and delivered it with minimal input from me, while I held down the sauropod side of our expanding empire of neural canal projects. She has the paper mostly written, too.

Oh, and we did get the smoking-gun images I wanted, of SMAs communicating with pneumatic spaces in the vertebrae via foramina in the neural canal. Often these foramina go up into the neural arch and spine, but in some cases – notably in pelicans and the occasional ratite – they go down into the centrum. So I now have no excuse for not getting back to the sauropod SMA paper (among many other things).

We’re making this all available because not only are we not afraid of getting scooped, we’re trying to get the word out. SMAs are phylogenetically widespread in birds and we know they were present in sauropods as well, so we should see some evidence of them in theropods and pterosaurs (because reasons). I made such a nuisance of myself at the recent Flugsaurier meeting, talking to everyone who would listen about SMAs, that Dave Hone went and found some pneumatic foramina in the neural canals of Pteranodon vertebrae during the conference – I suspect just to shut me up. That’ll be some kind of Hone-Atterholt-Wedel-and-some-others joint before long, too.

Anyway, point is, SMAs are cool, and you now have everything you need to go find them in more critters. Jessie and I are happy to collaborate if you’re interested – if nothing else, we have the background, lit review, and phylogenetic sampling down tight – but we don’t own SMAs, and we’ll be nothing but thrilled when your own reports start rolling in. Unexplored anatomical territory beckons, people. Let’s do this.

References

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Left lateral view

Have we ever posted decent photos of the Brachiosaurus altithorax caudals? Has anyone? I can’t remember either thing ever happening. When I need images of brachiosaur bits, including caudals, I usually go to Taylor (2009).

Taylor (2009: fig. 3)

Which is silly, not because Mike’s diagrams compiling old illustrations aren’t good – they definitely are – but because I’m sitting on a war chest of decent photos of the actual material. I am home sick with a sore throat today, and I can’t be arsed to (1) follow up on the “Down in Flames” post, (2) add anything thoughtful to the vertebral orientation discussion, or (3) crop or color-adjust these photos. You’re getting them just as they came out of my camera, from my trip to the Field Museum in 2012.

Here are the rest of the orthogonal views:

Right lateral view

 

Anterior view

 

Posterior view

 

Dorsal view of caudal 1

 

Dorsal view of caudal 2

And here’s a virtual walkaround using a series of oblique shots. Making a set like this is part of my standard practice now for important specimens during museum visits.

 

 

 

 

 

 

 

Now, I said up top that I wasn’t going to add anything thoughtful to the vertebral orientation discussion. I have thoughts on that, but I’m tired and hopped up on cold medicine and now ain’t the time. In lieu of blather, here are a couple of relevant photos.

 

I wanted to capture for my future self the pronounced non-orthogonality of the neural canal and centrum, so I rolled up a piece of paper and stuck it through the neural canal. I haven’t run the numbers, but in terms of “angle of the articular faces away from the neural canal”, these verts look like they’re right up there with my beloved Snowmass Haplocanthosaurus.

More on that next time, I reckon. In the meantime, all these photos are yours now (CC-BY, like everything on this site [that someone else hasn’t asserted copyright over]). Go have fun.

Reference

We all know that apatosaurines have big honkin’ cervical ribs (well, most of us know that). But did they also have unusually large neural spines?

The question occurred to me the other day when I was driving home from work. I was thinking about C10 of CM 3018, the holotype of Apatosaurus louisae, and I thought, “Man, that is a lot of neural spine right there.”

Why was I thinking about C10, particularly? I traced and also stacked Gilmore’s (1936) drawing for my 2002 paper with Kent Sanders, and recycled the trace for my 2007 prosauropod paper, and recycled the stack-o-C10s for my 2013 PeerJ paper with Mike. So for better or worse C10 is my mental shorthand for A. louisae, the same way that their respective C8s seem to capture the essence of Giraffatitan and Sauroposeidon.

I decided that the quick-and-dirty solution was to compare the vertebrae of A. louisae with those of Diplodocus carnegii, the default reference diplodocid, and see how they stacked up. With the cotyles scaled to the same vertical diameters, this is what we get for C9 and C10 of CM 3018 (lighter gray, background, traced from Gilmore 1936) vs CM 84/94 (darker gray, foreground, traced from Hatcher 1901):

The A. louisae verts are a hair taller, proportionally, than those of D. carnegii, but not by much. The difference is trivial compared to the differences in centrum length and cervical rib size.

So where did I get this apparently erroneous impression that Apatosaurus had giant neural spines? Maybe it’s not that the neural spines of apatosaurines in particular are so large, but rather than diplodocids of all types have large neural spines compared to non-diplodocids. Here are the same vertebrae compared for D. carnegii (dark gray, background) and Camarasaurus supremus (black, foreground, traced from Osborn and Mook 1921):

I deliberately picked the longest C9 in the AMNH collection, and the least-distorted C10. The first surprise for me was how well this C. supremus C9 hangs with D. carnegii in terms of proportions. That is one looooong Cam vert. In any other sauropod, it would probably be beautiful. But because it’s Camarasaurus it attained its length in the most lumpen possible way, with the diapophysis way up front, the neural spine apex way at the back, and in the middle just…more vertebra. Like a stretch limo made from a Ford Pinto, or Mike’s horrifying BOBA-horse.

Inevitable and entirely justified Cam-bashing aside, it’s striking how much smaller the whole neural arch-and-spine complex is in C. supremus than in D. carnegii. And remember that D. carnegii is itself a bit smaller than Apatosaurus, spine-wise. Is this maybe a diplodocoid-vs-macronarian thing, at least in the Morrison? Here’s the C10 stack with Brachiosaurus included, represented by BYU 12867 (which I think is probably a C10 based on both centrum proportions and neural spine shape – see Wedel et al. 2000b for details), and with labels added because it’s getting a little nuts:

I like this; it shows a lot. Here are some things to note:

  • The diplodocids don’t just have taller neural spines, their pre- and postzygapophyses are also higher than in the macronarians. That’s gotta mean something, right? All else being equal, putting the zygs farther from the intervertebral joints would reduce the flexibility of the neck. Maybe diplodocoids could get away with it because they had more cervicals, or maybe their necks were stiffened for some reason.
  • The zygs being set forward of their respective centrum ends in the macronarians really comes through here.
  • The Brachiosaurus vert isn’t that different from a stretched (and de-uglified) Cam vert, with a slightly higher neural spine to help support the longer neck. (Maybe this is why Cam inspires such visceral revulsion: it reads as a failed brachiosaur.)
  • This emphasizes the outlier status of Apatosaurus in the cervical rib department. It bears repeating: the cervical ribs of Camarasaurus are certainly wide, but they’re not nearly as massive or ventrally expanded as in apatosaurines.

So far, pretty interesting. I’d like to add Barosaurus and Haplocanthosaurus to round out the “big six” Morrison sauropods. I known Haplo has big, tall, almost apatosaurine neural spines (as shown above, with arrows highlighting the epipophyses), but for Baro I’d have to actually do the comparison to see where it falls out.

The idea of bringing in Barosaurus also forces the question, previously glossed over, of how legit it is to compare C10s of all these animals when their cervical counts differed. C. supremus is thought to have had 12 vertebrae in its neck, Brachiosaurus 13 (based on Giraffatitan), A. louisae and D. carnegii 15, and Barosaurus probably 16. It would be more informative to graph neural spine height divided by cotyle diameter along the column for all of these critters, plus Kaatedocus and Galeamopus. But that’s a lot of actual work, and as much fun as it sounds (really, I’d rather be doing that), I have summer teaching to prep for and field gear to wrangle. So I’ll have to revisit this stuff another time.

References

I know, I know — you never believed this day would come. And who could blame you? Nearly thirteen years after my 2005 SVPCA talkSweet Seventy-Five and Never Been Kissed, I am finally kicking the Archbishop descriptive work into gear. And I’m doing it in the open!

In the past, I’ve written my academic works in LibreOffice, submitted them for peer-review, and only allowed the world to see them after they’ve been revised, accepted and published. More recently, I’ve been using preprints to make my submitted drafts public before peer review. But there’s no compelling reason not to go more open than that, so I’ll be writing this paper out in the open, in a public GitHub repository than anyone can access. That also means anyone can file issues if they thing there’s something wrong or missing, and anyone can submit pull-requests if they have a correction to contribute.

I’ll be writing this paper in GitHub Flavoured Markdown so that it displays correctly right in the browser, and so that patches can be supported. That will make tables a bit more cumbersome, but it should be manageable.

Anyway, feel free to follow progress at https://github.com/MikeTaylor/palaeo-archbishop

The very very skeletal manuscript is at https://github.com/MikeTaylor/palaeo-archbishop/blob/master/archbishop-manuscript.md

Remember this broken Giraffatitan dorsal vertebra, which Janensch figured in 1950?

It is not only cracked in half, anteroposteriorly, it’s also unfused.

Here’s a better view of the broken face, more clearly showing that the neural canal is (a) much taller than wide – unlike all vertebrate spinal cords – and (b) almost entirely situated ventral to the neurocentral joint, getting close to the condition in the perverted Camarasaurus figured by Marsh.

Here’s a dorsal view, anterior to the top, with Mike’s distal forelimbs for scale.

Left lateral view.

Right lateral view – note the subtle asymmetries in the pneumatic foramen/camera. A little of that might be taphonomic distortion but I think much of it is real (and expected, most pneumatic systems produce asymmetries).

And postero-dorsal view, really showing the weird neural canal to good advantage. In this photo and in the pure dorsal view, you can see that the two platforms for the “neural arch” – which, as in the aforementioned Camarasaurus, is neither neural nor an arch – converge so closely as to leave only a paper-thin gap.

A few points arise. As explained in this post, it makes more sense to talk about the neurocentral joint migrating up or down relative to the neural canal, which is right where it always is, just dorsal to the articular faces of the centrum.

So far, in verts I’ve seen with “offset” neurocentral joints, the joint tends to migrate dorsally in dorsal vertebrae, putting the canal inside the developmental domain of the centrum (which now includes a partial or total arch in an architectural sense, even though the chunk of bone we normally call the neural arch develops as a separate bit) – as shown in the first post in this series. In sacral and caudal vertebrae, the situation is usually reversed, with the joint shifted down into what would normally be the centrum, and the canal then mostly or completely surrounded by the arch – as shown in the second post in the series. This post then doesn’t really add any new concepts, just a new example.

Crucially, we can only study this in the vertebrae of juveniles and subadults, because once the neurocentral joints are fused and remodeled, we usually can’t tell where the old joint surface was. So it’s like cervicodorsal and caudal dorsal pneumatic hiatuses, in that the feature of interest only exists for part of the ontogeny of the animal, and our sample size is therefore inherently limited. Not necessarily limited by material – most museums I’ve visited have a fair amount of juvenile and subadult material in the collections – but limited in published visibility, in that for many sauropods only the largest and most complete specimens have been monographically described.

So once again, the answer is simply to visit collections, look at lots of fossils, and stay alert for weird stuff – happily, a route that is open to everyone with a legitimate research interest.

Reference

  • Janensch, W. 1950. Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica (Suppl. 7) 3:27-93.

 

As part of a major spring cleaning operation that we started the first week of January, this week I opened the last two boxes left over from when we moved into our current house. One of them had a bundle of framed art. I knew most of what was in there before I opened the box, but I had somehow completely forgotten about this. I must have gotten it framed in late 90s, and it hung on the walls of our apartments in Norman and Santa Cruz. At some point it went into a box, and I forgot it even existed.

This is the first technical drawing I ever attempted of OMNH 53062, which would later become the holotype of Sauroposeidon. I drew it for my poster at the 1997 SVP meeting in Chicago, and it went on to become Figure 5 in my undergraduate thesis (which is preserved for posterity here). I’d do other, better drawings of the specimen in later years, but this one came first.

I know I’m biased, but that second vertebra in the preserved series, which I interpreted as C6 back when, will probably always be the most gorgeous natural object on the planet in my book. I don’t expect anyone else to feel the same. I worked on that specimen for three years – some of it seeped into my soul, and vice versa. Then again, I don’t care how jaded you are about long vertebrae, that one is still a pretty arresting sight.

For a much more recent take on the appearance of the Sauroposeidon vertebrae, see this post.

When I was nine, a copy of Don Glut’s The New Dinosaur Dictionary turned up in my local Waldenbooks. It wasn’t my first dinosaur book, by far – I’d been a dinosaurophile since the age of three. But The New Dinosaur Dictionary was different.

Up to that point, I had subsisted on a heavy diet of kids’ dino books and the occasional article in National Geographic and Ranger Rick. The kids’ books were aimed at kids and the magazine articles were pitched at an engagingly popular level. I didn’t understand every word, but they were clearly written for curious layfolk, not specialists.

A typical spread from The New Dinosaur Dictionary (Glut, 1982). The armored sauropod blew my young mind.

The New Dinosaur Dictionary was something else entirely. It had photos of actual dinosaur bones and illustrations of skeletons with cryptic captions like, “Skeleton of Daspletosaurus torosus. (After Russell)”. Okay, clearly this Russell cove was out there drawing dinosaur skeletons and this book had reproduced some of them. But nobody I knew talked like that, and the books I had access to up to that point held no comparable language.

The New Dinosaur Dictionary (Glut, 1982: p. 271)

Then there was stuff like this: “The so-called Von Hughenden sauropod restored as a brachiosaurid by Mark Hallett”. A chain of fascinating and pleasurable ideas detonated in my brain. “The so-called” – say what now? Nobody even knew what to call this thing? Somehow I had inadvertently sailed right to the edge of human knowledge of dinosaurs, and was peering out into taxa incognita. “Restored as a brachiosaurid” – so this was just one of several possible ways that the animal might have looked. Even the scientists weren’t sure. This was a far cry from the bland assurances and blithely patronizing tones of all my previous dinosaur books.

“By Mark Hallett.” I didn’t know who this Hallett guy was, but his art was all over the book, along with William Stout and some guy named Robert T. Bakker and a host of others who were exploding my conception of what paleo art could even be. Anyway, this Mark Hallett was someone to watch, not only because he got mentioned by name a lot, but because his art had a crisp quality that teetered on some hypercanny ridge between photorealism and scribbling. His sketches looked like they might just walk off the page.

In case that line about scribbling sounds dismissive: I have always preferred sketches by my favorite artists to their finished products. The polished works are frequently inhumanly good. They seem to have descended in a state of completed perfection from some divine realm, unattainable by mere mortals. Whereas sketches give us a look under the hood, and show how a good artist can conjure light, shadow, form, weight, and texture from a few pencil strokes. Put it this way: I am anatomist by temperament first, and by training and occupation second. Of course I want to see how things are put together.

The New Dinosaur Dictionary (Glut, 1982: p. 75)

Anyway, The New Dinosaur Dictionary was something completely new in my experience. It wasn’t aimed at kids and written as if by kids, like lots of kids’ books. It wasn’t even written by adults talking down (deliberately or inadvertently) to kids, or trying to reach a wide audience that might include kids. It was written by an adult, aiming at other adults. And it was admitting in plain language that we didn’t know everything yet, that there were lots of animals trembling on the outer threshold of scientific knowledge. I didn’t understand half of it – I was down in an ontogenetic trench, looking up as these packets of information exploded like fireworks over my head.

In Seeing In the Dark, the best book about why you should go out stargazing for yourself, Timothy Ferris writes about growing up on Florida’s Space Coast in the early 1960s, and watching the first generation of artificial satellites pass overhead:

I felt like an ancient lungfish contemplating the land from the sea. We could get up there.

That’s precisely the effect that The New Dinosaur Dictionary had on me: I could get up there. Maybe not immediately. But there were steps, bodies of knowledge that could be mastered piecemeal, and most of all, mysteries to be resolved. The book itself was like a sketch, showing how from isolated and broken bones and incomplete skeletons, scientists and artists reconstructed the world of the past, one hypothesis at a time. Now I take it for granted, because I’ve been behind the curtain for a couple of decades. But to my 9-year-old self, it was revolutionary.

This has all come roaring back because of something that came in the mail this week. Or rather, something that had been waiting in the mailroom for a while, that I finally picked up this week: a package from Mark Hallett, enclosing a copy of his 2018 dinosaur calendar. And also this:

 

An original sketch, which he gave to me as a Christmas present. The published version appears on one of the final pages of our book, where we discuss the boundaries between the known – the emerging synthesis of sauropod biology that we hoped to bring to a broader audience by writing the book in the first place – and the unknown – the enduring mysteries that Mark and I think will drive research in sauropod paleobiology for the next few decades. Presented without a caption or commentary, the sketch embodies sauropods as we see them: emerging from uncertainty and ignorance one hard-won line at a time, with ever-increasing solidity.

Thank you, Mark, sincerely. That sketch, what it evokes, both for me now and for my inner 9-year-old – you couldn’t have chosen a better gift. And I couldn’t be happier. Except perhaps to someday learn that our book exploded in the mind of a curious kid the way that The New Dinosaur Dictionary did for me 34 years ago, a time that now seems as distant and romantic as the primeval forests of the Mesozoic.