Regular readers will remember that we followed up our 1VPC talk about what it means for a vertebra to be horizontal by writing it up as a paper, and doing it in the open. That manuscripts is now complete, and published as a preprint (Taylor and Wedel 2019).

Taylor and Wedel (2018: Figure 5). Haplocanthosaurus sp. MWC 8028, caudal vertebra ?3, in cross section, showing medial aspect of left side, cranial to the right, in three orientations. A. In “articular surfaces vertical” orientation (method 2 of this paper). The green line joins the dorsal and ventral margins of the caudal articular surface, and is oriented vertically; the red line joins the dorsal and ventral margins of the cranial articular surface, and is nearly but not exactly vertical, instead inclining slightly forwards. B. In “neural canal horizontal” orientation (method 3 of this paper). The green line joins the cranial and caudal margins of the floor of the neural canal, and is oriented horizontally; the red line joins the cranial and caudal margins of the roof of the neural canal, and is close to horizontal but inclined upwards. C. In “similarity in articulation” orientation (method 4 of this paper). Two copies of the same vertebra, held in the same orientation, are articulated optimally, then the group is rotated until the two are level. The green line connects the uppermost point of the prezygapophyseal rami of the two copies, and is horizontal; but a horizontal line could join the two copies of any point. It happens that for this vertebra methods 3 and 4 (parts B and C of this illustration) give very similar results, but this is accidental.

The preprint has all the illustrations and their captions at the back of the PDF. If you prefer to have them inline in the text, where they’re referenced — and who wouldn’t? — you can download a better version of the manuscript from the GitHub archive.

By the way, you may have noticed that what started our written in Markdown has mutated into an MS-Word document. Why? Well, because journals won’t accept submissions in Markdown. It eas a tedious and error-prone job to convert the Markdown into MS-Word, and not one I am keen to repeat. For this reason, I think I am unlikely to use Markdown again for papers.

References

  • Taylor, Michael P., and Mathew J. Wedel. 2019. What do we mean by the directions “cranial” and “caudal” on a vertebra? PeerJ PrePrints 7:e27437v2. doi:10.7287/peerj.preprints.27437v2

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My friend and frequent collaborator Jessie Atterholt has her office in the next building over from mine. When you walk in, you see something that looks approximately like this. Not exactly like this, because I took these photos in February and she’s changed a few things (and I’m rubbish about getting stuff posted in a timely fashion).

The last time I showed an office full of amazing stuff like this, it was Peter Dodson’s. It will come as no surprise that Jessie was Peter’s student at UPenn before she went to Berkeley for her PhD.

The far case holds mostly books and skulls. Dr. A has her own plastination setup for making preserved organs and organisms, and the snake on the second shelf here is one that she prepped herself. One side of the snake still has the skin on, the other half has been skinned to show the muscles. This is crunch week for me so I don’t have time to ID all of the stuff, but alert readers should have no problem spotting some digitally-resurrected Haplocanthosaurus bits.

Mostly skulls on the middle rack. The sirenian skull on the second shelf and the cave bear on the fourth are both casts, but almost everything else is real bone. The bighorn sheep on the middle shelf is a natural mummy.

Here’s a close-up of the top shelf. Other than some 3D-printed human skull bones sitting in front of the brain slice on the left, everything here is real bone, including the lion, baboon, and human skulls, and the giraffe cervicals winding across the top. Jessie’s been collecting since she was a kid and the African megafauna are gifts from a globe-trotting family friend.

The upper shelves here have quite a few of Jessie’s plastinated specimens, both whole organisms and things like hearts and kidneys from various critters.

A close-up of some of Jessie’s coolest anatomical preparations. In back is an internal cast of the lungs and bronchial tree of a cat. The baby rattlesnake died after eating a proportionally gigantic lizard — I was dumb and forgot to flip the snake over to show the lizard inside, plastinated along with its predator. The ground squirrel on the right is another half-fleshed, half-skinned plastinate, and the mouse up front is a classic dissection presentation, preserved forever through plastination.

I’ve heard it said that the difference between a collector and a hoarder is curation. As someone who definitely lurks more on the hoarder end of that spectrum (to paraphrase Dave Barry, if you could see my office you’d be blinded or driven insane), I’m pretty darned jealous of both the breadth of Jessie’s collection, and the skill and taste with which it is displayed. She’s featured some of these specimens on her Instagram, which I strongly recommend.

In a word, amazingly. After 6 days (counting public galleries last Sunday), 4300 photos, 55 videos, dozens of pages of notes, and hundreds of measurements, we’re tired, happy, and buzzing with new observations and ideas.

We caught up with some old friends. Here Mike is showing an entirely normal and healthy level of excitement about meeting CM 584, a specimen of Camarasaurus from Sheep Creek, Wyoming. You may recognize this view of these dorsals from Figure 9 in our 2013 PeerJ paper.

We spent an inordinate amount of time in the public galleries, checking out the mounted skeletons of Apatosaurus and Diplodocus (and Gilmore’s baby Cam, and the two tyrannosaurs, and, and…).

I had planned a trip to the Carnegie primarily to have another look at the Haplocanthosaurus holotypes, CM 572 and CM 879. I was also happy for the chance to photograph and measure these vertebrae, CM 36034, which I think have never been formally described or referred to Haplocanthosaurus. As far as I know, other than a brief mention in McIntosh (1981) they have not been published on at all. I’m planning on changing that in the near future, as part of the larger Haplocanthosaurus project that now bestrides my career like a colossus.

The real colossus of the trip was CM 555, which we’ve already blogged about a couple of times. Just laying out all of the vertebrae and logging serial changes was hugely useful.

Incidentally, in previous posts and some upcoming videos, we’ve referred to this specimen as Brontosaurus excelsus, because McIntosh (1981) said that it might belong to Apatosaurus excelsus. I was so busy measuring and photographing stuff that it wasn’t until Friday that I realized that McIntosh made that call because CM 555 is from the same locality as CM 563, now UWGM 15556, which was long thought to be Apatosaurus excelsus but which is now (i.e., Tschopp et al. 2015) referred to Brontosaurus parvus. So CM 555 is almost certainly B. parvus, not B. excelsus, and in comparing the specimen to Gilmore’s (1936) plates of CM 563, Mike and I thought they were a very good match.

Finding the tray of CM 555 cervical ribs was a huge moment. It added a ton of work to our to-do lists. First we had to match the ribs to their vertebrae. Most of them had field numbers, but some didn’t. Quite a few were broken and needed to be repaired – that’s what I’m doing in the above photo. Then they all had to be measured and photographed.

It’s amazing how useful it was to be able to reassociate the vertebrae with their ribs. We only did the full reassembly for c6, in part because it was the most complete and perfect of all of the vertebrae, and in part because we simply ran out of time. As Mike observed in his recent post, it was stunning how the apatosaurine identity of the specimen snapped into focus as soon as we could see a whole cervical vertebra put back together with all of its bits.

We also measured and photographed the limb bones, including the bite marks on the radius (above, in two pieces) and ulna (below, one piece). Those will of course go into the description.

And there WILL BE a description. We measured and photographed every element, shot video of many of them, and took pages and pages of notes. Describing even an incomplete sauropod skeleton is a big job, so don’t expect that paper this year, but it will be along in due course. CM 555 may not be the most complete Brontosaurus skeleton in the world, but our ambition is to make it the best-documented.

In the meantime, we hopefully left things better documented than they had been. All of the separate bits of the CM 555 vertebrae – the centra, arches, and cervicals ribs – now have the cervical numbers written on in archival ink (with permission from collections manager Amy Henrici, of course), so the next person to look at them can match them up with less faffing about.

We have people to thank. We had lunch almost every day at Sushi Fuku at 120 Oakland Avenue, just a couple of blocks down Forbes Avenue from the museum. We got to know the manager, Jeremy Gest, and his staff, who were unfailingly friendly and helpful, and who kept us running on top-notch food. So we kept going back. If you find yourself in Pittsburgh, check ’em out. Make time for a sandwich at Primanti Bros., too.

We owe a huge thanks to Calder Dudgeon, who took us up to the skylight catwalk to get the dorsal-view photos of the mounted skeletons (see this post), and especially to Dan Pickering, who moved pallets in collections using the forklift, and moved the lift around the mounted skeletons on Tuesday. Despite about a million ad hoc requests, he never lost patience with us, and in fact he found lots of little ways to help us get our observations and data faster and with less hassle.

Our biggest thanks go to collections manager Amy Henrici, who made the whole week just run smoothly for us. Whatever we needed, she’d find. If we needed something moved, or if we needed to get someplace, she’d figure out how to do it. She was always interested, always cheerful, always helpful. I usually can’t sustain that level of positivity for a whole day, much less a week. So thank you, Amy, sincerely. You have a world-class collection. We’re glad it’s in such good hands.

What’s next? We’ll be posting about stuff we saw and learned in the Carnegie Museum for a long time, probably. And we have manuscripts to get cranking on, some of which were already gestating and just needed the Carnegie visit to push to completion. As always, watch this space.

References

If you were curious about the Wedel et al. presentation on the Snowmass Haplocanthosaurus at the 1st Palaeo Virtual Congress but didn’t attend the event, it is now preserved for posterity and freely available to the world as a PeerJ Preprint (as promised). Here’s the link.

I’ll have much more to say about this going forward, but for now here are slides 20 and 21 on the intervertebral joint spaces. This is obviously just the same vert cloned three times and articulated with itself. With the digital rearticulation of the reconstructed and retrodeformed caudal series still in progress, we cloned caudal 3, the only vertebra that preserves both sets of zygapophyses, to get a rough estimate of the sizes and shapes of the soft tissues that filled the intervertebral spaces and neural canal.

The reconstructed intervertebral discs (in blue) are very crude and diagrammatic. The reason I’m putting these particular slides up is to get the cited references out in the open on the blog, to start correcting the misapprehension that all non-mammalian amniotes have exclusively synovial intervertebral joints (see the discussion in the comments on this post). In the list below I’m including Banerji (1957), which is not cited in the presentation but which I did cite in that comment thread; it’s an important source and at least for now it is a free download. These refs are just the tip of a very big iceberg. One of my goals for 2019 is to do a series of posts reviewing the extensive literature on amphiarthrodial (fibrocartilaginous) intervertebral joints in living lepidosaurs and birds. Stay tuned!

And please go have a look at the presentation if you are at all interested or curious. As we said in the next to last slide, “this research is ongoing, and we welcome your input. If there are facts or hypotheses we haven’t considered but should, please let us know!”

References

The 1st Palaeontological Virtual Congress is underway now, and will run through December 15. Mike and I have two presentations up:

“What do we mean by the directions ‘cranial’ and ‘caudal’ on a vertebra?” by Mike and me, which consists of a video Mike made presenting a slide show that he put together. The presentation sums up our thinking following the series of vertebral orientation posts here earlier this summer and fall, which are all available here.

“Reconstructing an unusual specimen of Haplocanthosaurus using a blend of physical and digital techniques” by me and a gang of WesternU-based collaborators, including Jessie Atterholt and Thierra Nalley, both of whom you saw in our recent pig-hemisecting adventures. Almost everything I’ve written on this blog about Haplocanthosaurus in 2018 was part of the run-up to this presentation (except, somewhat ironically, the post about pneumaticity), which also includes quite a bit that I haven’t put on the blog yet. So even if you follow SV-POW!, the 1PVC slideshow should have plenty of stuff you haven’t seen yet.

IF you can see it–you have to be a registered 1PVC ‘attendee’ to log in to the site and see the presentations. So probably you are either already registered and this post is old news, or not registered and this post seems useless. Why would I bother telling you about stuff you can’t see?

The answer is that neither Mike or I intend for our work to disappear when 1PVC comes to an end on December 15. Both of us are planning to put our abstracts and slide decks up as PeerJ Preprints, which is our default move for conference presentations these days (e.g., this, this, and this). I believe Mike is also going to post his video to YouTube. So the work will not only live on after the congress is over, it will jump to a much broader audience. We’re looking forward to letting everyone see what we’ve been up to, and I’m sure we’ll have some more things to say here when that happens.

So, er, go see our stuff if you’re a 1PVC attendee, and if you’re not, hang in there, we’ll have that stuff out to you in a few days. UPDATE: The Haplo presentation is up now (link).

Here’s D10 and the sacrum of Diplodocus AMNH 516 in left lateral and ventral view, from Osborn (1904: fig. 3). Note how the big lateral pneumatic foramina, here labeled ‘pleurocoelia’, start out up at the top of the centrum in D10 and kind of pinch out up there, seemingly entirely absent by S3 (although there is a suspicious-looking shadowed spot above and behind the sacral rib stump labeled ‘r3’). Then on S4 and S5 the big foramina are back, but now they’re low on the centrum, ventral to the sacral ribs. In ventral view, the foramina on D10, S1, and S2 aren’t visible–they’re both over the curve of the centrum, and in the case of S1 and S2, obscured by the sacral ribs. But in S4 and S5, the big lateral foramina are visible in ventral view.

I’ve been interested in a while in this seeming hand-off in centrum pneumatization from dorsolateral, which prevails in the dorsal vertebrae, to ventrolateral, which prevails in the posterior sacral and caudal vertebrae. Almost all sauropod dorsals have the pneumatic foramina quite high on the centrum, sometimes even encroaching on the neural arch. But if sauropod caudals have pneumatic fossae or foramina on the centrum, they’re usually quite low, and almost always below the caudal rib or transverse process (there may also be pneumatic fossae on the neural arch and spine)–for evidence, see Wedel and Taylor (2013b). To me this implies two different sets of diverticula.

I think that in part because sometimes you get both sets of diverticula acting on a single vert. Here’s the centrum of sacral 4 of Haplocanthosaurus CM 879 in right dorsolateral view; anterior is to the right.

Here’s the same thing annotated (yeah, it does look a little like an Ent who is alarmed because his left eye has been overgrown by a huge nasal tumor). This vert has two sets of pneumatic features on the centrum: a big lateral fossa below the sacral rib articulation, presumably homologous with the same feature in S4 of the Diplodocus above; and a smaller dorsolateral fossa above and behind sacral rib articulation.

Unfortunately, CM 879 doesn’t tell us much about how these two sets of diverticula might have changed along the column. The centra of S1-S3 were not found, S5 lacks both sets of fossae, the first caudal has fossae both on the centrum, below the caudal rib, and low on the arch, and the second and subsequent caudals lack both sets of fossae. (I wrote a LOT more about pneumaticity in this individual in my 2009 air sacs paper, which is linked below.)

Working out how these diverticula change serially is a tractable problem. 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. As far as I know, that’s never been done, but feel free to correct me if I’ve missed something. I’m neck deep in other stuff, so if someone wants that project, have at it. (If you happen to look into this, I’d be grateful for a heads up, so we don’t run over each other if I do get a yen to investigate further myself.)

References

WOW! I knew I was dragging a bit on getting around to this vertebral orientation problem, but I didn’t realize a whole month had passed. Yikes. Thanks to everyone who has commented so far, and thanks to Mike for getting the ball rolling on this. Previous posts in this series are here and here.

First up, this may seem like a pointlessly picky thing to even worry about. Can’t we just orient the vertebrae in whichever way feels the most natural, or is easiest? Do we have to think about this?

The alarmingly 3D pelvis of the mounted brontosaur at the AMNH. Note that sauropod pubes are usually illustrated lying flat, so what usually passes for ‘lateral’ view would be roughly from the point of view of the animal’s knee.

I think we do. For sauropods, vertebrae are usually oriented for illustration purposes in one of two ways. The first is however they sit most easily on their pallets. This is similar to the problem Mike and I found for ‘lateral’ views of sauropod pelvic elements when were on our AMNH/Yale trip in 2012. In an articulated skeleton, the pubes and ischia usually lean inward by 30-45 degrees from their articulations with the ilia, so they can meet on the midline, but when people illustrate the “lateral view” of a sauropod pubis or ischium, it’s often the ventro-lateral aspect that is face-up when the element is lying on a shelf or a pallet. Photographic lateral does not equal biological lateral for those elements. Similarly, if I’m trying to answer biological questions about vertebrae (see below), I need to know something about their orientation in the body, not just how they sit comfortably on a pallet.

The other way that vertebrae are commonly oriented is according to what we might call the “visual long axis” of the centrum—so for example, dorsoventrally tall but craniocaudally short proximal caudals get oriented with the centrum ‘upright’, whereas dorsoventrally short but craniocaudally long distal caudals get oriented with the centrum ‘horizontal’, even if they’re in the same tail and doing so makes the neural canals or articular faces be oriented inconsistently down the column. (I’m not going to name names, because it seems mean to pick on people for something I just started thinking about myself, but if you go plow through a bunch of sauropod descriptions, you’ll see what I’m talking about.)

Are there biological questions where this matters? You bet! There are some questions that we can’t answer unless we have the vertebrae correctly oriented first. One that comes to mind is measuring the cross-sectional area of the neural canal, which Emily Giffin did a lot of back in the 90s. Especially for the Snowmass Haplocanthosaurus, what counts as the cross-sectional area of the neural canal depends on whether we are looking at the verts orthogonal to their articular faces, or in alignment with the course of the canal. I think the latter is pretty obviously the way to go if we are measuring the cross-sectional area of the canal to try and infer the diameter of the spinal cord—we’d want to see the canal the same way the cord ‘sees’ it as it passes through—but it’s less obvious if we’re measuring, say, the surface area of the articular face of the vertebra to figure out, say, cartilage stress. It doesn’t seem unreasonable to me that we might want to define a ‘neural axis’ for dealing with spinal-cord-related questions, and a ‘biomechanical axis’ for dealing with articulation-related questions.

Caudal 3 of the Snowmass Haplocanthosaurus, hemisected 3D model.

With all that in mind, here are some points.

To me, asking “how do we know if a vertebra is horizontal” is an odd phrasing of the problem, because “horizontal” doesn’t have any biological meaning. I think it makes more sense to couch the question as, “how do we define cranial and caudal for a vertebra?” Normally both the articular surfaces and the neural canal are “aimed” head- and tail-wards, so the question doesn’t come up. Our question is, how do we deal with vertebrae for which the articular surfaces and neural canal give different answers?

For example. Varanus komodoensis caudal.

(And by the way, I’m totally fine using “anterior” and “posterior” for quadrupedal animals like sauropods. I don’t think it causes any confusion, any more than people are confused by “superior” and “inferior” for human vertebrae. But precisely because we’re angling for a universal solution here, I think using “cranial” and “caudal” makes the most sense, just this once. That said, when I made the image above, I used anterior and posterior, and I’m too lazy now to change it.)

I think if we couch the question as “how do we define cranial and caudal”, it sets up a different set of possible answers than Mike proposed in the first post in this series: (1) define cranial and caudal according to the neural canal, and then describe the articular surfaces as inclined or tilted relative to that axis; (2) vice versa—realizing that using the articular surfaces to define the anatomical directions may admit a range of possible solutions, which might resurrect some of the array of possible methods from our first-draft abstract; (3) define cranial and caudal along the long axis of the centrum, which is potentially different from either of the above; (4) we can imagine a range of other possibilities, like “use the zygs” or “make the transverse processes horizontal” (both of which are subsets of Mike’s method C) but I don’t think most of those other possibilities are sufficiently compelling to be worthy of lengthy discussion.

IF we accept “neural canal”, “articular surfaces”, and “centrum long axis” as our strongest contenders, I think it makes most sense to go with the neural canal, for several reasons:

  • In a causative sense, the neural tube/spinal cord does define the cranial/caudal axis for the developing skeleton. EDIT: Actually, that’s a bit backwards. It’s the notochord, which is later replaced by the vertebral column, that induces the formation of the brain and spinal cord from the neural plate. But it’s still true that the vertebrae form around the spinal cord, so it’s not wrong to talk about the spinal cord as a defining bit of soft tissue for the developing vertebrae to accommodate.
  • The neural canal works equally well for isolated vertebrae and for articulated series. Regardless of how the vertebral column is oriented in life, the neural canal is relatively smooth—it may bend, but it doesn’t kink. So if we line up a series of vertebrae so that their neural canals are aligned, we’re probably pretty close to the actual alignment in life, even before we look at the articular surfaces or zygs.
  • The articulated tails of Opisthocoelicaudia and big varanids show that sometimes the articular surfaces simply are tilted to anything that we might reasonably consider to be the cranio-caudal axis or long axis of the vertebra. In those cases, the articular surfaces aren’t orthogonal to horizontal OR to cranio-caudal. So I think articular surfaces are ruled out because they break down in the kinds of edge cases that led us to ask the question in the first places.

Opistocoelicaudia caudals 6-8, stereopair, Borsuk-Bialynicka (1977:plate 5).

“Orient vertebrae, isolated or in series, so that their neural canals define the cranio-caudal axis” may seem like kind of a ‘duh’ conclusion (if you accept that it’s correct; you may not!), but as discussed up top, often vertebrae from a single individual are oriented inconsistently in descriptive works, and orientation does actually matter for answering some kinds of questions. So regardless of which conclusion we settle on, there is a need to sort out this problem.

That’s where I’m at with my thinking. A lot of this has been percolating in my hindbrain over the last few weeks—I figured out most of this while I was writing this very post. Is it compelling? Am I talking nonsense? Let me know in the comments.